In defence of the entity of Macaronesia as a biogeographical region.

Since its coinage ca. 1850 AD by Philip Barker Webb, the biogeographical region of Macaronesia, consisting of the North Atlantic volcanic archipelagos of the Azores, Madeira with the tiny Selvagens, the Canaries and Cabo Verde, and for some authors different continental coastal strips, has been under dispute. Herein, after a brief introduction on the terminology and purpose of regionalism, we recover the origins of the Macaronesia name, concept and geographical adscription, as well as its biogeographical implications and how different authors have positioned themselves, using distinct terrestrial or marine floristic and/or faunistic taxa distributions and relationships for accepting or rejecting the existence of this biogeographical region. Four main issues related to Macaronesia are thoroughly discussed: (i) its independence from the Mediterranean phytogeographical region; (ii) discrepancies according to different taxa analysed; (iii) its geographical limits and the role of the continental enclave(s), and, (iv) the validity of the phytogeographical region level. We conclude that Macaronesia has its own identity and a sound phytogeographical foundation, and that this is mainly based on three different floristic components that are shared by the Macaronesian core (Madeira and the Canaries) and the outermost archipelagos (Azores and Cabo Verde). These floristic components are: (i) the Palaeotropical-Tethyan Geoflora, formerly much more widely distributed in Europe and North Africa and currently restricted to the three northern archipelagos (the Azores, Madeira and the Canaries); (ii) the African Rand Flora, still extant in the coastal margins of Africa and Arabia, and present in the southern archipelagos (Madeira, the Canaries and Cabo Verde), and (iii) the Macaronesian neoendemic floristic component, represented in all the archipelagos, a result of allopatric diversification promoted by isolation of Mediterranean ancestors that manage to colonize Central Macaronesia and, from there, the outer archipelagos. Finally, a differentiating floristic component recently colonized the different archipelagos from the nearest continental coast, providing them with different biogeographic flavours.

A multitaxa approach to biodiversity inventory in Matela protected area (Terceira, Azores, Portugal).

Background: This manuscript is the first contribution of the project, "Matela - uma ilha de biodiversidade" ("Matela - an island of biodiversity"), that aims to restore the native vegetation within the Azorean Protected Area of the Terceira Island Nature Park known as the "Protected Area for the Management of Habitats or Species of Matela" (TER08), situated on Terceira Island, the Azores Archipelago, Portugal. This small fragment of native forest, positioned at a low-medium altitude (300-400 m a.s.l.), is facing some conservation impacts as a consequence of the spread of different invasive exotic plant species, mainly Pittosporumundulatum, Rubusulmifolius and Hedychiumgardnerianum. The database we present encompasses diverse taxonomic groups, including bryophytes, vascular plants, arthropods, birds and mammals. It is derived from intensive sampling campaigns conducted in 2022, but some data from a previous vascular plant survey in 2015 were also included. The objective of this study was to provide an updated inventory of bryophytes, vascular plants, arthropods, birds and mammals within this protected area. In this way we are providing the reference conditions necessary for the monitoring of the impacts of the current ongoing restoration efforts within the project "Matela - an island of biodiversity". Whenever feasible, the present inventory is juxtaposed with historical data from previous surveys conducted in Matela. New information: In the realm of bryophytes, our analysis revealed the presence of 75 taxa, comprising 44 mosses and 32 liverworts. Amongst these, 71 were indigenous, while three remained indeterminate and one, Campylopusintroflexus, was identified as invasive. A comparison with previous historical data revealed a decrease in species richness, which was partially counterbalanced by the discovery of 23 new recorded species in the area.Regarding vascular plants, we distinguished 54 species, comprising 28 indigenous and 26 introduced taxa. Almost 80% of the inventoried species (n = 43) were newly documented in Matela.The study of arthropods encompassed a total of 103 taxa. Within the realm of soil arthropods, we documented eight indigenous and 25 introduced taxa, witnessing the disappearance of endemic species alongside a substantial increase in introduced ones between 2002 and 2022. Canopy arthropods, totalling 36 indigenous and 18 introduced taxa, exhibited few changes when compared with data from 2002. SLAM traps captured 24 indigenous and 15 introduced arthropod taxa and no historical data are available for comparison.As for avian species, we noted 12 indigenous birds and one introduced species, confirming the presence of most of the historical recorded native species.The mammalian census revealed eight introduced species, setting new precedents for Matela, alongside the identification of one endemic species: the Azorean endemic bat Nyctalusazoreum.

Vascular plant taxa occurrences in exotic woodland and in natural and production forests on the Islands of São Miguel, Terceira and Pico (Azores).

Background: The data presented here originated from field expeditions carried out between 2017 and 2018, within the framework of Forest-Eco2 project: "Towards an Ecological and Economic valorisation of the Azorean Forest". The project aimed to quantify the ecological value of the Azorean forests, including carbon accumulation and to design and propose measures that could further enhance forest sustainability. For that, 90 forest plots were sampled on three Azores islands - São Miguel, Terceira and Pico - equally distributed into natural forest, exotic woodland and production forest. The aim of this report is to further expand knowledge on biodiversity trends enclosed in the different forest types present in the Azores, by providing a list of the occurrences of the 105 different vascular plant taxa together with a brief characterisation of their origin and life-form. New information: We provide an inventory of indigenous and non-indigenous vascular plant taxa from 90 forest stands. A total of 105 taxa were identified and registered, belonging to 60 families, 91 genera, 101 species and four subspecies. A total of 35% of the taxa were endemic, 27% native and 38% non-indigenous, including 19% of invasive taxa. Endangered and vulnerable taxa were registered, including Elaphoglossumhirtum (Sw.) C.Chr., Lactucawatsoniana Trel. and others which were considered by the authors a priority for conservation (e.g. Arceuthobiumazoricum Wiens & Hawksw., Bellisazorica Hochst. ex Seub., Saniculaazorica Guthnick ex Seub., Platantheramicrantha (Hochst. ex Seub.) Schltr.). Our records provide detailed and updated knowledge of Azorean Forest flora and highlight the role of natural forests as indigenous plant diversity hotspots and exotic woodland as a source of invasive taxa within the Archipelago.

Assessing the effects of climate change on arthropod abundance in Azorean pastures: PASTURCLIM project's baseline monitoring data.

Background: The data we present are part of the project PASTURCLIM (Impact of climate change on pasture's productivity and nutritional composition in the Azores). The project aims to assess the consequences of climate change (e.g. temperature increase) on the grass production and its quality for forage, as well as to assess changes in the arthropod communities associated with the Azorean intensive pastures. An in situ experiment was set up using Open Top Chambers (OTCs), in order to simulate an increasing of temperature (average of +1.2ºC) on pastures. In this contribution, we present the data relative to the arthropod sampling. New information: We provide an inventory of all arthropods recorded inside OTCs and in control plots in three intensively managed pastures dominated by grasses in Terceira Island (Azores): two of them dominated by ryegrass, Loliummultiflorum Lam. (Poaceae), located respectively at 186 m and 301 m above sea level; and one field dominated by common velvetgrass, Holcuslanatus L. (Poaceae), located at an altitude of 385 m.A total of 41351 specimens were collected. Organisms collected belong to four classes, 15 orders, 60 families and 171 species/morphospecies (including 34 taxa identified only at order, family or genus level). Therefore, for only 137 taxa, we have a scientific name associated (n = 38918). A total of 75% of the species (n = 129 species) are considered introduced (including all the species with indeterminate colonisation status that are possibly also exotic species (n = 7622)), representing 71% of the total abundance (n = 29664 specimens). A total of 19% of the species (n = 33 species) are considered native non-endemic representing 28% of the total abundance (n = 11608 specimens). Only one endemic species was sampled, the wolf spider Pardosaacorensis Simon, 1883 (1% of the species), representing 0.2% of the total abundance (n = 79 specimens). Spiders (5056 specimens) and beetles (18310 specimens) were the dominant taxa representing, respectively, 20 and 78 morphospecies.Since the main aim of this study was to have a better knowledge on arthropod communities present in Azorean pastures under a simulated temperature increase, the principal novelty of this paper is the contribution with distribution and abundance data to a baseline knowledge on the future consequences of climate changes on arthropod communities in Azorean pastures.

Taxonomic, structural diversity and carbon stocks in a gradient of island forests.

Assessment of forest ecosystems and their services is seen as a key action for the advancement of biodiversity objectives, and to inform the development and implementation of related policies and planning. The Azorean forest is important for timber production, the protection of soil and water resources, and for its recreational and aesthetic value. However, its role in carbon accumulation has not been fully addressed. We assessed plant diversity, forest structure and carbon stocks in a gradient of three forest types (Natural Forest-NF; Exotic Woodland-EW and Production Forest-PF) in three of the Azores islands. We used biodiversity indices and found that NF harbored the highest plant diversity levels and PF the lowest. Diversity levels were lower for structural than for taxonomic data, particularly for PF. The highest tree carbon stock was found at EW in one of the islands, while PF consistently exhibited relatively high tree carbon stocks in the three islands. The largest soil carbon stocks were found at EW, while leaf litter carbon stocks were higher at PF. We concluded that NF play a fundamental role as plant diversity hotspots but have lower relevance as carbon stocks what might be associated with montane environmental conditions. PFs provide economic assets and act as carbon sinks, while EWs play a major role as carbon sinks in soil, but also at tree level in the oldest forests.

Arthropods and other biota associated with the Azorean trees and shrubs: Laurusazorica (Seub) Franco (Magnoliophyta, Magnoliopsida, Laurales, Lauraceae).

This study explores the composition and structure of species communities associated with the native Azorean tree species Laurusazorica (Seub) Franco (Magnoliophyta, Magnoliopsida, Laurales, Lauraceae). Communities were sampled in six Islands covering the occidental (Flores), central (Faial, Pico, Terceira) and eastern (São Miguel, Santa Maria) groups of Azores Archipelago during the BALA project, using standardised sampling protocols for surveying canopy arthropod fauna. In addition, the study characterises the distribution of species regarding their colonisation status and feeding modes and, finally, compares communities of different Islands. Ninety-four arthropod species totalling 10,313 specimens were collected on L.azorica. The Arthropod community was dominated by Hemiptera species, most of them being herbivores. Endemic and native species showed a very high abundance representing about 94% of the total species abundance. However, despite introduced species being represented by few individuals (6% of the total abundance), their diversity was remarkable (28 species and no significant difference with diversity found in endemic and native species communities). Analysis of rarity patterns revealed a stable community of endemic species (alpha gambin SAD model approaching a log-normal shape), intermediate stable community of native species (alpha SAD gambin model approaching a poisson log-normal) and a less stable community of introduced species (alpha SAD gambin model approaching a log-series shape). A dissimilarity analysis revealed high similarity between communities of Terceira and Pico and high dissimilarity between Flores and Faial communities. We observed a clear individualisation of the different islands when considering endemic species, whereas we observed high overlap when considering native and introduced species groups. Canopy community distribution confirms the results obtained in a previous study which suggest the stability of native and endemic arthropods species communities over introduced species community in native forests fragments. Arthropod species were richer than bryophytes, lichens and vascular plants species. We found that L.azorica serve as the substrate for very few vascular plants species (four epiphytes species), which were present in all Islands, except Elaphoglossumsemicylindricum, which does not occur in Santa Maria. L.azorica shelters a significant number of bryophytes and lichens species. Thirty-two lichens and 92 bryophyte species, including 57 liverworts and 35 mosses, are referred to this phorophyte. Five bryophyte species, all Azorean endemics, are considered Endangered by IUCN Criteria. L.azorica harbours a poor community of epiphyte vascular plant species and all of them were ferns, but the community of bryophytes and lichens are not negligible although very low compared to the community found on other previously studied Azorean trees, the Azorean cedar Juniperusbrevifolia. The present study shows that most islands present particular species distribution patterns without geographical correlation and that conservation programmes should be adapted to each Island. The study, therefore, calls for a specialisation of conservation programmes for each of the Islands.

Biota from the coastal wetlands of Praia da Vitória (Terceira, Azores, Portugal): Part 4 - Vascular plants.

BACKGROUND: The data presented here come from field observations, carried out between 2014 and 2017, as part of a LIFE research project aiming to preserve and restore three coastal wetlands of Praia da Vitória (Terceira Island, Azores, Portugal) (LIFE-CWR). A total of 23 vascular plant species surveys were carried out in three sites: one for each semester in Paul da Praia da Vitória (PPV) and Paul da Pedreira do Cabo da Praia (PPCP); one for each semester (except in 2014) in Paul do Belo Jardim (PBJ). The main objectives were to determine the plant richness of the three sites and to monitor yearly variation on species composition. NEW INFORMATION: A total of 107 taxa, belonging to 50 families, were observed, many of which are new records for the area, especially in PBJ and PPCP, where 78 and 92% of species records were new. A few very rare species in the Azores were recorded in these coastal wetlands, namely Lotus creticus, Bolboschoenus maritimus, Juncus maritimus and Polygonum maritimum.

The colonization history of Juniperus brevifolia (Cupressaceae) in the Azores Islands.

BACKGROUND: A central aim of island biogeography is to understand the colonization history of insular species using current distributions, fossil records and genetic diversity. Here, we analyze five plastid DNA regions of the endangered Juniperus brevifolia, which is endemic to the Azores archipelago. METHODOLOGY/PRINCIPAL FINDINGS: The phylogeny of the section Juniperus and the phylogeographic analyses of J. brevifolia based on the coalescence theory of allele (plastid) diversity suggest that: (1) a single introduction event likely occurred from Europe; (2) genetic diversification and inter-island dispersal postdated the emergence of the oldest island (Santa Maria, 8.12 Ma); (3) the genetic differentiation found in populations on the islands with higher age and smaller distance to the continent is significantly higher than that on the younger, more remote ones; (4) the high number of haplotypes observed (16), and the widespread distribution of the most frequent and ancestral ones across the archipelago, are indicating early diversification, demographic expansion, and recurrent dispersal. In contrast, restriction of six of the seven derived haplotypes to single islands is construed as reflecting significant isolation time prior to colonization. CONCLUSIONS/SIGNIFICANCE: Our phylogeographic reconstruction points to the sequence of island emergence as the key factor to explain the distribution of plastid DNA variation. The reproductive traits of this juniper species (anemophily, ornithochory, multi-seeded cones), together with its broad ecological range, appear to be largely responsible for recurrent inter-island colonization of ancestral haplotypes. In contrast, certain delay in colonization of new haplotypes may reflect intraspecific habitat competition on islands where this juniper was already present.

Genetic variability and differentiation among populations of the Azorean endemic gymnosperm Juniperus brevifolia: baseline information for a conservation and restoration perspective.

The Azorean endemic gymnosperm Juniperus brevifolia (Seub.) Antoine is a top priority species for conservation in Macaronesia, based on its ecological significance in natural plant communities. To evaluate genetic variability and differentiation among J. brevifolia populations from the Azorean archipelago, we studied 15 ISSR and 15 RAPD markers in 178 individuals from 18 populations. The average number of polymorphic bands per population was 65 for both ISSR and RAPD. The majority of genetic variability was found within populations and among populations within islands, and this partitioning of variability was confirmed by AMOVA. The large majority of population pairwise F(ST) values were above 0.3 and below 0.6. The degree of population genetic differentiation in J. brevifolia was relatively high compared with other species, including Juniperus spp. The genetic differentiation among populations suggests that provenance should be considered when formulating augmentation or reintroduction strategies.