Abstract Abstract
Background
During the LIFE-CWR project "Ecological Restoration and Conservation of Praia da Vitória Coastal Wet Green Infrastructures", there was the opportunity to undertake a systematic record of bryophytes at Paul da Praia da Vitória (PPV), Paul do Belo Jardim (PBJ) and Paul da Pedreira do Cabo da Praia (PPCP), three coastal wetland areas of Praia da Vitória (Terceira, Azores, Portugal). The objective of the study was to perform a biodiversity assessment, comparing the three sites at two different moments, before and after the implementation of several conservation measures. This project also contributed to improve the knowledge of Azorean bryophyte diversity at both local and regional scales, including the recording of two new taxa for the Azores and three new taxa for Terceira Island.
New information
This paper reports the results of the first extensive survey of bryophyes of the three coastal wetland areas of Praia da Vitória (Terceira Island, Azores, Portugal). The identification of a total of 504 samples, 240 collected in 2013 and 2016 (before the LIFE-CWR intervention) and 265 in 2017 and 2018 (after the intervention), resulted in a list of 58 species of bryophytes (one hornwort, eight liverworts and 48 mosses). These include two new records for the Azores (Bryum klinggraeffii, Ptychostomum bornholmense), three new records for Terceira Island (Bryum tenuisetum, Dicranella howei, Trichostomum crispulum) and at least 15 new records for the municipality of Praia da Vitória (e.g., Cephaloziella hampeana). Some species that had not been recorded on the island since 1937 (e.g., Fissidens crispus) were collected during this study, as well as a Macaronesian endemic liverwort (Radula wichurae), an Iberian-Macaronesian liverwort (Frullania azorica) and a moss species with European distribution (Ptychomitrium nigrescens). From the recorded species, only one moss (Leptophascum leptophyllum), of subtropical origin, is considered invasive in Europe.
Keywords: bryophytes, mosses, liverworts, hornworts, coastal wetlands, bryophyte surveys, standardised sampling, Terceira Island (Azores)
Introduction
Bryophytes are small plants without vascular tissues, directly depending on immediate environmental conditions. Most species are poikilohydric, i.e., lacking the ability (structural or functional) to maintain and/or regulate water content to achieve homeostasis of cells and tissue. Because they are so dependant on their immediate environment, they respond quickly to environmental change, which makes them good bioindicators of changes in land use, precipitation regime, temperature, salinity and pollution.
The Azores archipelago is well-known for its rich bryoflora (480 species and subspecies) (Gabriel et al. 2010), which may be related to the high humidity and mild temperatures, influenced by the Atlantic Ocean, and scarce pollution sources.
Although the coastal areas of the islands are among their most deteriorated habitats, mainly due to urbanization pressure, some interesting, though fragmented, ecosystems remain at lower elevations in the Azores. The three coastal wetlands of Praia da Vitória municipality are a case in point, well worth restoration and habitat protection. The areas, studied during the LIFE-CWR Project – Paul da Praia da Vitória (PPV), Paul do Belo Jardim (PBJ) and Paul da Pedreira do Cabo da Praia (PPCP) – are best known for their birds (Barcelos et al. 2015; Dias 2018; Goulart et al. 2019); however, they harbour other important and interesting biological groups, such as molluscs and arthropods (Martins and Borges 2019), plants and lichens (Elias et al. 2019).
The coastal areas of the Azorean Islands are not thoroughly studied, since bryologists tend to focus on the rich natural forests of the archipelago (e.g., Aranda et al. 2011) and the bryophytic flora of these coastal wetlands had never been systematically sampled.
This is the second contribution in a series of papers (Borges et al. 2018) intending to characterize the biota of the three areas.
General description
Purpose
The main aim of this work was to inventory the bryophyte species present in Paul da Praia da Vitória (PPV), Paul do Belo Jardim (PBJ) and Paul da Pedreira do Cabo da Praia (PPCP), three neighbouring areas focused on by the restoration project LIFE-CWR, in order to improve knowledge on the regional distribution of bryophytes (mosses, liverworts, hornworts) and set a baseline for future research in the area.
Project description
Title
Inventory of bryophytes in three coastal wetlands of Terceira Island (Azores)
Personnel
The inventory was conducted during the years of 2013, 2016, 2017 and 2018 under the responsibility of Rosalina Gabriel, with the participation of Javier Diaz Castillo (2013), César Pimentel and Mariana Reis Brito (2016, 2017). Sampling dates and collectors are listed in Table 1. Species identifications were performed by César Pimentel under the supervision of Rosalina Gabriel (2016 and 2017), and David Claro (2013) under the supervision of Cecília Sérgio. The identification of some challenging samples was performed by Manuela Sim-Sim and Cecília Sérgio.
Table 1.
Dates of collection and collectors of bryophytes from the three wetlands of the county of Praia da Vitória (PPV - Paul da Praia da Vitória; PBJ - Paul do Belo Jardim; PPCP - Paul da Pedreira do Cabo da Praia).
| Area | Year | Sampling date | Transects | Latitude | Longitude | Collectors |
| PPV | 2016 | 31-03-2016 | 1 | 38,73534 | -27,06042 | César Pimentel |
| 01-04-2016 | 2 | 38,73449 | -27,05833 | César Pimentel & Mariana R. Brito | ||
| 28-06-2016 | 3 | 38,73590 | -27,06027 | César Pimentel & Mariana R. Brito | ||
| 28-06-2016 | 4 | 38,73443 | -27,05944 | César Pimentel & Mariana R. Brito | ||
| 2017 | 12-06-2017 | 1A | 38,73443 | -27,05944 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |
| 2A | 38,73534 | -27,06042 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 3A | 38,73449 | -27,05833 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 4A | 38,73590 | -27,06027 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| PBJ | 2013 | 18-10-2013 | 1A | 38,71337 | -27,06080 | Javier Diaz Castillo |
| 28-10-2013 | 1B | 38,71298 | -27,06102 | Javier Diaz Castillo | ||
| 2A | 38,71389 | -27,06119 | Javier Diaz Castillo | |||
| 01-11-2013 | 2B | 38,71317 | -27,06123 | Javier Diaz Castillo | ||
| 2017 | 13-06-2017 | 1 | 38,71355 | -27,06182 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |
| 2 | 38,71350 | -27,06107 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 3 | 38,71333 | -27,06113 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| PPCP | 2016 | 24-02-2016 | 1 | 38,70367 | -27,04556 | Rosalina Gabriel, César Pimentel & Mariana R. Brito |
| 2 | 38,70327 | -27,04553 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 13-05-2016 | 3 | 38,70440 | -27,04513 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | ||
| 4 | 38,70267 | -27,04801 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 2017 | 14-06-2017 | 1A | 38,70367 | -27,04556 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |
| 2A | 38,70327 | -27,04553 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 3A | 38,70440 | -27,04513 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 4A | 38,70267 | -27,04801 | Rosalina Gabriel, César Pimentel & Mariana R. Brito | |||
| 2018 | 01-09-2018 | na | 38,70306 | -27,04745 | Rosalina Gabriel & Paulo A.V. Borges |
Study area description
Terceira Island (area: 400.6 km²; elevation: 1,021.14 m) is one of the nine islands of the Azores archipelago, located in the North Atlantic, roughly at 38°43′49″N, 27°19′10″W (Forjaz 2004). The climate in the Azores is temperate oceanic, with regular and abundant rainfall, high levels of relative humidity and persistent western winds, mainly during the winter and autumn seasons (Azevedo et al. 2004).
Terceira Island is known for the presence of some very important native forest areas at high elevation (e.g., Gabriel and Bates 2005). However, few natural areas remain at lower elevations, notably in Praia da Vitória county. Three coastal wetland areas were studied in this project: Paul da Praia da Vitória (PPV) (Figs 1, 2), Paul do Belo Jardim (PBJ) (Figs 3, 4) and Paul da Pedreira do Cabo da Praia (PPCP) (Figs 5, 6).
Figure 1.
General aspect of Paul da Praia da Vitória with its islands and surrounding urban area (Photo by Paulo A.V. Borges).
Figure 2.
Detail of the recently created "islands" of Juncus acutus in Paul da Praia da Vitória (Photo by Paulo A.V. Borges).
Figure 3.
Paul do Belo Jardim' s dune area (Photo by Paulo A.V. Borges).
Figure 4.
Juncus maritimus growing in the Paul do Belo Jardim area (Photo by Paulo A.V. Borges).
Figure 5.
General view of Paul da Pedreira do Cabo da Praia, at low tide (Photo by Paulo A.V. Borges).
Figure 6.
Detail of the margin of Paul da Pedreira do Cabo da Praia during high tide (Photo by Paulo A.V. Borges).
Potentially, the plant cover of Praia da Vitória would include Erica-Morella coastal woodlands (cf. Elias et al. 2016). However, apart from some native shrubs of Morella faya, still found in PPV and PPCP, the main native species found in the area include species typical of humid zones, namely Juncus (J. acutus, J. maritimus, J. effusus) and Ruppia maritima (Elias et al. 2019). Presently, most of the area is covered with exotic and invasive species, with the giant cane (Arundo donax), being especially abundant in PBJ, and the sticky snakeroot (Ageratina adenophora) in PPCP. Both these species are included in the first quartile of invasive species in the three archipelagos of Macaronesia (Silva et al. 2008). The bryophytic component of the flora had previously not been systematically studied in the area.
Design description
In each wetland, a network of three (PBJ) or four (PPV, PPCP) transects (160 m × 2 m or 300 m × 2 m), was set and sampled every 20 m (ocasionally every 10 m), in quadrats/sampling points of 4 m2; each quadrat was searched for bryophytes. Whenever possible, a maximum of six samples or microplots (10 cm × 5 cm) were collected: three replicates from soil and three replicates from rock. Bryophyte samples were brought to the laboratory for identification and herborization at the Herbarium of the University of Azores (AZU) – section Bryophytes.
In 2018, after the project's completion, the areas continued to be visited and in one of those visits a new species for the Paul da Pedreira do Cabo da Praia location was collected.
Funding
This study was financed by the project LIFE+ (LIFE12 BIO/PT/000110: Ecological Restoration and Conservation Infrastructure Green Wet Coast Praia da Vitória) (2013–2018).
Sampling methods
Study extent
This study coverered a small coastal area, extending from PPV (to the North) to PPCP (to the South), with an extent of 3.58 km.
Sampling description
In each site, bryophytes were sampled using standardised methods, during one or two visits in 2013, 2016 and 2017, respectively (Table 1). Within each transect, a quadrat with an area of 4 m2 was delimited at intervals of 10 or 20 meters. Each sampling point (quadrat) was carefully examined to collect three samples (replicates or microplots) for each of the available substrates (soil, rock); microplots were randomly selected from areas colonized by bryophytes. The area collected in each replica (microplot) was 50 cm2 (10 cm × 5 cm). The maximum number of samples obtained per sampling point was six (three replicates of bryophytes growing on soil and three of bryophytes growing on rock), but many sampling points did not contain bryophytes, especially those located in areas periodically flooded with brackish waters. For each microplot, some ecological variables were also measured (e.g., insulation, water availability) using ordinal scales adapted from Gabriel and Bates (2005), and some soil or rock was recovered for pH measurement.
Quality control
The correct identification of the sampled taxa is crucial in an inventory. Keys and floras were used to identify the species, and their coverage (in %) was also estimated for each microplot in the laboratory. The main floras used for the identification of liverworts were by Paton (1999) and Casas et al. (2009), whereas for mosses Smith (2004), Casas et al. (2006) and different volumes of "Flora Briofítica Ibérica" (Guerra 2018) were used. Taxonomic keys provided by Schumacker and Váňa (2005) and field guides (Atherton et al. 2010, Llimona et al. 2004) were also checked. Some important internet databases were consulted, namely the Azorean Biodiversity Portal and TROPICOS for taxonomic data and BBS Field Guide online pages, Bildatlas der Moose Deutschlands and Swissbryophytes for morphological and ecological data. Nomenclature mostly follows Gabriel et al. (2010) and Ros et al. 2013.
Samples were mostly examined by CP and DC, and their identification was supervised by RG and CS.
Geographic coverage
Description
Praia da Vitória municipality, Terceira Island, Azores archipelago, Macaronesia, Portugal.
Coordinates
38º42'09''N and 38°44'12''N Latitude; 27º03'46''W and 27°02'39''W Longitude.
Taxonomic coverage
Description
Bryophytes, including Division Anthocerotophyta, Division Bryophyta, and Division Marchantiophyta.
Temporal coverage
Notes
The main sampling was performed in 2013, 2016 and 2017; a single sample was collected in 2018.
Usage rights
Use license
Open Data Commons Attribution License
IP rights notes
Additional information on this study may also be requested from the first author.
Data resources
Data package title
LIFE_CWR_TER_Bryophytes
Resource link
Alternative identifiers
Number of data sets
1
Data set 1.
Data set name
Bryophytes from Praia da Vitória.
Data format
Darwin Core Archive.
Number of columns
56
Download URL
Data format version
1.
Description
In this data table, we include all records for which a taxonomic identification of the species was possible. 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 (DwC-A), which is a standardised format for sharing biodiversity data as a set of one or more data tables. The core data table contains 188 events. One extension data table also exists. 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 downloadin the downloads section. The versions table lists other versions of the resource that have been made publicly available and allows tracking changes made to the resource over time.
Data set 1.
| Column label | Column description |
|---|---|
| Table Event | The sub-table with events |
| id | Identifier of the events, unique for the dataset |
| type | Type of the record, as defined by the Public Core standard |
| license | Reference to the license under which the record is published |
| institutionID | The identity of the institution publishing the data |
| institutionCode | The code of the institution publishing the data |
| datasetName | Name of the dataset |
| eventID | Identifier of the events, unique for the dataset |
| samplingProtocol | The sampling protocol used to capture the species |
| samplingEffort | The amount of time of each sampling |
| eventDate | The date-time or interval during which an Event occurred. For occurrences, this is the date-time when the event was recorded. |
| startDayOfYear | The earliest ordinal day of the year on which the Event occurred |
| habitat | The habitat for an Event |
| continent | The name of the continent in which the Location occurs |
| islandGroup | The name of the island group in which the Location occurs |
| island | The name of the island on or near which the Location occurs |
| country | The name of the country or major administrative unit in which the Location occurs |
| countryCode | The standard code for the country in which the Location occurs |
| municipality | The full, unabbreviated name of the next smaller administrative region than county (city, municipality, etc.) in which the Location occurs |
| locality | The specific description of the place |
| verbatimCoordinates | Original coordinates recorded |
| decimalLatitude | Approximate centre point decimal latitude of the field site in GPS coordinates |
| decimalLongitude | Approximate centre point decimal longitude of the field site in GPS coordinates |
| Table Occurrences | The sub-table with occurrence data |
| id | Identifier of the events, unique for the dataset |
| license | Reference to the license under which the record is published |
| institutionID | The identity of the institution publishing the data |
| institutionCode | The code of the institution publishing the data |
| collectionCode | The code of the collection where the specimens are conserved |
| datasetName | Name of the dataset |
| basisOfRecord | The nature of the data record |
| dynamicProperties | A list of additional measurements, facts, characteristics, or assertions about the record. Meant to provide a mechanism for structured content |
| occurrenceID | Identifier of the record, coded as a global unique identifier |
| occurrenceRemarks | Remarks on the occurrence substracte from where the specimens were captured |
| recordNumber | An identifier given to the Occurrence at the time it was recorded |
| recordedBy | A list (concatenated and separated) of names of people, groups, or organizations responsible for recording the original Occurrence |
| organismQuantity | A number or enumeration value for the quantity of organisms |
| organismQuantityType | The unit of the identification of the organisms |
| establishmentMeans | The process of establishment of the species in the location, using a controlled vocabulary: 'native non-endemic', 'introduced', 'endemic' |
| disposition | The current state of a specimen with respect to the collection identified in collectionCode or collectionID |
| eventID | Identifier of the events, unique for the dataset |
| fieldNumber | An identifier given to the event in the field |
| minimumElevationInMeters | Minimum elevation in metres |
| identifiedBy | Name of the person who made the identification |
| dateIdentified | Date on which the record was identified |
| scientificName | Complete scientific name including author |
| kingdom | Kingdom name |
| phylum | Phylum name |
| class | Class name |
| order | Order name |
| family | Family name |
| genus | Genus name |
| specificEpithet | Specific epithet |
| infraspecificEpithet | Infraspecific epithet |
| taxonRank | Lowest taxonomic rank of the record |
| scientificNameAuthorship | The authorship information for the scientificName formatted according to the conventions of the applicable nomenclaturalCode |
Additional information
The identification of the samples (242 before the LIFE-CWR intervention [2013, 2016], 261 after it [2017]) resulted in a set of 57 species of bryophytes, including one hornwort, eight liverwort species (Table 2) and 48 moss species (Table 3), representing about 80% of the bryophyte species present in the three sampled areas, according to the first-order Jackknife estimator (Table 4) (Gabriel 2018).
Table 2.
List of hornworts (Division Anthocerotophyta) and liverworts (Division Marchantiophyta) identified in the three coastal wetlands of Praia da Vitória, Terceira Island, Azores, Portugal (PPV – Paul da Praia da Vitória; PBJ – Paul do Belo Jardim; PPCP – Paul da Pedreira do Cabo da Praia), indicating their Class, Order and Family as well as the number of samples obtained for each species during LIFE-CWR fieldwork.
| Class | Order | Family | Species/Subspecies | PPV | PBJ | PPCP |
| Anthocerotopsida | Notothyladales | Notothyladaceae | Phaeoceros laevis (L.) Prosk. | 1 | 2 | |
| Jungermanniopsida | Fossombroniales | Fossombroniaceae | Fossombronia caespitiformis De Not. ex Rabenh. subsp. multispira (Schiffn.) J. R. Bray et D. C. Cargill | 9 | ||
| Jungermanniopsida | Jungermanniales | Cephaloziellaceae | Cephaloziella hampeana (Nees) Schiffn. | 2 | ||
| Jungermanniopsida | Porellales | Frullaniaceae | Frullania azorica Sim-Sim et al. | 2 | 8 | |
| Jungermanniopsida | Porellales | Lejeuneaceae | Marchesinia mackaii (Hook.) Gray | 1 | ||
| Jungermanniopsida | Porellales | Radulaceae | Radula lindenbergiana Gottsche ex C. Hartman | 1 | ||
| Jungermanniopsida | Porellales | Radulaceae | Radula wichurae Steph. | 2 | ||
| Marchantiopsida | Lunulariales | Lunulariaceae | Lunularia cruciata (L.) Dumort ex. Lindb. | 1 | 8 | 7 |
| Marchantiopsida | Marchantiales | Ricciaceae | Riccia huebeneriana Lindenb. | 1 |
Table 3.
List of mosses (Division Bryophyta) identified in the three coastal wetlands of Praia da Vitória, Terceira Island, Azores, Portugal (PPV – Paul da Praia da Vitória; PBJ – Paul do Belo Jardim; PPCP – Paul da Pedreira do Cabo da Praia), indicating their Class, Order and Family as well as the number of samples obtained for each species during LIFE-CWR fieldwork.
| Class | Order | Family | Species/Subspecies | PPV | PBJ | PPCP |
| Bryopsida | Bartramiales | Bartramiaceae | Philonotis marchica (Hedw.) Brid. | 7 | ||
| Bryopsida | Bartramiales | Bartramiaceae | Philonotis rigida Brid. | 1 | 2 | |
| Bryopsida | Bryales | Bryaceae | Anomobryum julaceum (P. Gaerth., B. Mey. et Scherb.) Schimp. | 45 | ||
| Bryopsida | Bryales | Bryaceae | Bryum argenteum Hedw. | 9 | ||
| Bryopsida | Bryales | Bryaceae | Bryum canariense Brid. | 4 | ||
| Bryopsida | Bryales | Bryaceae | Bryum klinggraeffii Schimp. | 4 | ||
| Bryopsida | Bryales | Bryaceae | Bryum ruderale Crundw. et Nyholm | 5 | 1 | |
| Bryopsida | Bryales | Bryaceae | Bryum subapiculatum Hampe | 1 | 14 | |
| Bryopsida | Bryales | Bryaceae | Bryum tenuisetum Limpr. | 1 | 16 | |
| Bryopsida | Bryales | Bryaceae | Ptychostomum capillare (Hedw.) D. T. Holyoak et N. Pedersen | 36 | 84 | |
| Bryopsida | Bryales | Bryaceae | Ptychostomum dichotomum Hedw. | 1 | ||
| Bryopsida | Bryales | Bryaceae | Ptychostomum bornholmense (Wink. & R.Ruthe) Holyoak & N.Pedersen | 7 | ||
| Bryopsida | Bryales | Bryaceae | Ptychostomum pseudotriquetrum (Hedw.) J. R. Spence et H. P. Ramsay ex D. T. Holyoak et N. Pedersen | 1 | ||
| Bryopsida | Bryales | Bryaceae | Ptychostomum rubens (Mitt.) D. T. Holyoak et N. Pedersen | 12 | 6 | 2 |
| Bryopsida | Dicranales | Dicranaceae | Dicranella howei Renauld et Cardot | 1 | 1 | |
| Bryopsida | Dicranales | Ditrichaceae | Ceratodon purpureus (Hedw.) Brid. subsp. purpureus | 4 | ||
| Bryopsida | Dicranales | Fissidentaceae | Fissidens crispus Mont. | 61 | 7 | 12 |
| Bryopsida | Dicranales | Fissidentaceae | Fissidens viridulus (Sw. ex anon.) Wahlenb. | 1 | ||
| Bryopsida | Dicranales | Leucobryaceae | Campylopus pilifer Brid. | 31 | ||
| Bryopsida | Grimmiales | Grimmiaceae | Grimmia lisae De Not. | 3 | 11 | 57 |
| Bryopsida | Grimmiales | Ptychomitriaceae | Ptychomitrium nigrescens (Kunze) Wijk et Marg. | 1 | ||
| Bryopsida | Hypnales | Brachytheciaceae | Brachytheciastrum velutinum (Hedw.) Ignatov et Huttunen | 7 | 1 | 4 |
| Bryopsida | Hypnales | Brachytheciaceae | Brachythecium mildeanum (Schimp.) Milde | 1 | ||
| Bryopsida | Hypnales | Brachytheciaceae | Brachythecium rutabulum (Hedw.) Schimp. | 5 | 1 | 12 |
| Bryopsida | Hypnales | Brachytheciaceae | Brachythecium plumosum (Hedw.) Schimp. | 1 | ||
| Bryopsida | Hypnales | Brachytheciaceae | Kindbergia praelonga (Hedw.) Ochyra | 7 | 3 | 10 |
| Bryopsida | Hypnales | Brachytheciaceae | Oxyrrhynchium hians (Hedw.) Loeske | 1 | 1 | |
| Bryopsida | Hypnales | Brachytheciaceae | Oxyrrhynchium speciosum (Brid.) Warnst. | 1 | ||
| Bryopsida | Hypnales | Brachytheciaceae | Rhynchostegiella litorea (De Not.) Limpr. | 3 | ||
| Bryopsida | Hypnales | Brachytheciaceae | Rhynchostegium confertum (Dicks.) Schimp. | 17 | 6 | 19 |
| Bryopsida | Hypnales | Brachytheciaceae | Rhynchostegium megapolitanum (F. Weber et D. Mohr.) Schimp. | 1 | ||
| Bryopsida | Hypnales | Hypnaceae | Hypnum cupressiforme Hedw. var. cupressiforme | 15 | ||
| Bryopsida | Hypnales | Leucodontaceae | Leucodon sciuroides (Hedw.) Schwägr. | 1 | ||
| Bryopsida | Pottiales | Pottiaceae | Barbula convoluta Hedw. | 1 | 8 | 12 |
| Bryopsida | Pottiales | Pottiaceae | Barbula unguiculata Hedw. | 1 | ||
| Bryopsida | Pottiales | Pottiaceae | Didymodon australasiae (Hook. & Grev.) R.H. Zander | 6 | ||
| Bryopsida | Pottiales | Pottiaceae | Didymodon sicculus M.J. Cano, Ros, García-Zam. & J. Guerra | 11 | ||
| Bryopsida | Pottiales | Pottiaceae | Didymodon tophaceus (Brid.) Lisa | 1 | ||
| Bryopsida | Pottiales | Pottiaceae | Didymodon umbrosus (Müll. Hal.) R.H. Zander | 5 | 3 | 10 |
| Bryopsida | Pottiales | Pottiaceae | Leptophascum leptophyllum (Müll. Hal.) J. Guerra et M. J. Cano | 8 | 1 | |
| Bryopsida | Pottiales | Pottiaceae | Pseudocrossidium hornschuchianum (Schultz) R. H. Zander | 1 | ||
| Bryopsida | Pottiales | Pottiaceae | Tortella flavovirens (Bruch.) Broth. | 3 | 10 | 23 |
| Bryopsida | Pottiales | Pottiaceae | Tortula muralis Hedw. | 3 | 3 | 2 |
| Bryopsida | Pottiales | Pottiaceae | Tortula solmsii (Schimp.) Limpr. | 2 | 1 | |
| Bryopsida | Pottiales | Pottiaceae | Tortula truncata (Hedw.) Mitt. | 1 | ||
| Bryopsida | Pottiales | Pottiaceae | Trichostomum brachydontium Bruch | 14 | 12 | 119 |
| Bryopsida | Pottiales | Pottiaceae | Trichostomum crispulum Bruch | 2 | 15 | |
| Bryopsida | Pottiales | Pottiaceae | Weissia controversa Hedw. | 3 | ||
| Polytrichopsida | Polytrichales | Polytrichaceae | Polytrichum piliferum Hedw. | 1 |
Table 4.
Some statistical data from the collection of bryophytes in Praia da Vitória wetlands per year of collection (Number of samples; Observed richness or number of species (S); Number of estimated richness according to the first order Jackknife estimator; Percentage of completeness, i.e., ratio between the number of estimated species and the number of observed of species).
| 2013|2016 | 2017 | ||
| PPV | Number of samples | 44 | 83 |
| Observed richness (S) | 14 | 17 | |
| Estimated richness | 17,91 | 22,93 | |
| % Completeness | 78,17 | 74,14 | |
| PBJ | Number of samples | 42 | 58 |
| Observed richness (S) | 14 | 24 | |
| Estimated richness | 18,88 | 32,84 | |
| % Completeness | 74,15 | 73,08 | |
| PPCP | Number of samples | 156 | 120 |
| Observed richness (S) | 33 | 28 | |
| Estimated richness | 43,93 | 32,96 | |
| % Completeness | 75,12 | 84,95 | |
| TOTAL | Number of samples | 242 | 261 |
| Observed richness (S) | 45 | 42 | |
| Estimated richness | 57,95 | 52,96 | |
| % Completeness | 77,65 | 79,31 |
Comparison between years (before and after CWR intervention): The main interventions performed by the LIFE-CWR project in the three coastal areas included the removal of garbage and litter from PPCP, the opening of a small lagoon in PBJ and the connection of PPV to the sea.
The number of species varied slightly before and after the interventions, but the level of completeness is acceptable, higher than 75%, for both sampling periods (Table 4). The highest value of bryophyte species richness was observed in Paul da Pedreira do Cabo da Praia, probably due to the availability of a higher proportion of rocky substrata, while the lowest richness value was observed in Paul da Praia da Vitória.
A Ward's dissimilarity analysis performed with the diversity of species found at the three studied wetlands shows a remarkable homogeinity of results between the studied years (Fig. 7). Thus, the LIFE-CWR restoration interventions, especially focused on the improvement the bird habitat and water flow, did not hinder the conservation of bryophytes.
Figure 7.
Dendogram showing the result of a Ward's dissimilarity analysis including the bryophytes from the three wetlands of the municipality of Praia da Vitória (Terceira Island, Azores) before (2013, 2016) and after (2017) the interventions made by the LIFE-CWR project.
Main biogeographic distribution of the species: Most species found in the three studied wetlands have a broad biogeographic distribution, generally circumpolar and European, showing temperate climatic characteristics. Although most of the collected species are common in the Azores, three species are classified as Rare by IUCN (Grimmia lisae, Tortula solmsii and Riccia huebeneriana) (Dierssen 2001), and a single moss species (Leptophascum leptophyllum), of subtropical origin, is considered invasive in Europe. This species is widespread in humanized areas and may commonly be found on the sidewalks of some cities (Blockeel et al. 2014).
Noteworthy species: Among the observed species, two represent new records for the Azores, Bryum klinggraeffii (Ellis et al. 2016) and Ptychostomum bornholmense (Ellis et al. 2018). There are also three new records for Terceira Island (Bryum tenuisetum, Dicranella howei and Trichostomum crispulum) and at least 15 new recordsfor the municipality of Praia da Vitória, including the moss Leucodon sciuroides, a species previously known only from Monte Brasil (Angra do Heroísmo, Terceira Island) (Fontinha and Sérgio 1995) and which has been declining in the United Kingdom (Blockeel et al. 2014), and the liverwort Cephaloziella hampeana, also known from a single location on Terceira Island (Algar do Carvão) (Crundwell et al. 2013).
Some species that had not been recorded on Terceira Island since 1937 (e.g., Fissidens crispus) (Gabriel et al. 2011) were found on the wetlands, which may be explained by a lack of fieldwork at low elevations on the island (Aranda et al. 2011).
A Macaronesian endemic liverwort (Radula wichurae) and an Iberian-Macaronesian liverwort (Frullania azorica; Fig. 8) were found growing on rocks in the different wetlands. Actually, Praia da Vitória county, parish of Cabo da Praia, represents the classical locality of Frullania azorica, the place from where the species was originally collected and described (Sim-Sim et al. 1995). This species is frequently found in the area, sometimes forming extensive colonies on exposed rocks near the ocean. However, in this study, it was not identified in Paul da Praia da Vitória, possibly because there are not many rocks available for colonization.
Figure 8.
The liverwort Frullania azorica was described from material collected in the Praia da Vitória area (Photo by Rosalina Gabriel).
The acrocarpic moss species Ptychomitrium nigrescens, endemic to Europe and Macaronesia (Macaronesia, Portugal and France), was also reported from Paul da Pedreira do Cabo da Praia (PPCP), where boulders and large rocks are available for colonization.
Further details related to the LIFE-CWR project can be found in the book by Brian Morton, Elisabete Nogueira and António Frias Martins (Morton et al. 2019) and in the report by RG (Gabriel 2018).
Acknowledgements
We would like to acknowledge the financial support for field and lab work provided by the project LIFE CWR – Ecological Restoration and Conservation of Praia da Vitória Coastal Wet Green Infrastructure (2013-2018; Life12 bio7pt/000110). Open access was funded by FEDER (85%) and by Azorean Public funds (15%), through Operational Program Azores 2020 under the project AZORESBIOPORTAL – PORBIOTA (ACORES-01-0145-FEDER-000072).
We are deeply grateful to the leader of the LIFE CWR project, Eng.ª Elisabete Nogueira, for her visionary leadership on this project over the years, and to Dr. Rui Figueira and Tainan Messina, GBIF - Portugal, for the creation of the Darwin Core Archive.
We would also like to thank to the editor of BDJ, Yasen Mutafchiev, and the reviewers of our work, Michele Aleffi, Juana González-Mancebo, Niels Klazenga, Robert Mesibov and Tatyana Shubina, for their thoughtful feedback on an earlier version of the manuscript, which has greatly enhanced the final publication.
Author contributions
RG conceived the project. RG and PAVB conceived and drafted the manuscript. RG, CMMP, JDC and MRB collected the data. RG, CMMP, DC, CS and MSS identified the species. RG organised the final database. All authors revised and contributed to the final text.
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