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"Macaronesia"
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Island woodiness underpins accelerated disparification in plant radiations
The evolution of secondary (insular) woodiness and the rapid disparification of plant growth forms associated with island radiations show intriguing parallels between oceanic islands and tropical alpine sky islands. However, the evolutionary significance of these phenomena remains poorly understood and the focus of debate.
We explore the evolutionary dynamics of species diversification and trait disparification across evolutionary radiations in contrasting island systems compared with their nonisland relatives. We estimate rates of species diversification, growth form evolution and phenotypic space saturation for the classical oceanic island plant radiations – the Hawaiian silverswords and Macaronesian Echium – and the well-studied sky island radiations of Lupinus and Hypericum in the Andes.
We show that secondary woodiness is associated with dispersal to islands and with accelerated rates of species diversification, accelerated disparification of plant growth forms and occupancy of greater phenotypic trait space for island clades than their nonisland relatives, on both oceanic and sky islands.
We conclude that secondary woodiness is a prerequisite that could act as a key innovation, manifest as the potential to occupy greater trait space, for plant radiations on island systems in general, further emphasizing the importance of combinations of clade-specific traits and ecological opportunities in driving adaptive radiations.
Journal Article
Anthropogenic transitions from forested to human-dominated landscapes in southern Macaronesia
The extinction of iconic species such as the dodo and the deforestation of Easter Island are emblematic of the transformative impact of human colonization of many oceanic islands, especially those in the tropics and subtropics. Yet, the interaction of prehistoric and colonial-era colonists with the forests and forest resources they encountered can be complex, varies between islands, and remains poorly understood. Long-term ecological records (e.g., fossil pollen) provide the means to understand these human impacts in relation to natural change and variability pre- and postcolonization. Here we analyze paleoecological archives in forested landscapes of the Canary Islands and Cabo Verde, first colonized approximately 2,400 to 2,000 and 490 y ago, respectively. We demonstrate sensitivity to regional climate change prior to human colonization, followed by divergent but gradual impacts of early human settlement. These contrast with more rapid transformation in the colonial era, associated with significant increases in anthropogenic pressures. In the Canary Islands, at least two native tree taxa became extinct and lowland thermophilous woodlands were largely converted to agricultural land, yet relictual subtropical laurel forests persisted with limited incursion of nonnative species. In Cabo Verde, in contrast, thermophilous woodlands were depleted and substituted by open landscapes and introduced woodlands. Differences between these two archipelagos reflect the changing cultural practices and societal interactions with forests and illustrate the importance of long-term data series in understanding the human footprint on island ecosystems, information that will be critically important for current and future forest restoration and conservation management practices in these two biodiversity hotspots.
Journal Article
Shifts in grasses diversity patterns between two contrasting 40‐year climate periods in tropical dry islands
Grasses are one of the most successful and dispersed plant families worldwide and their environmental and economic values are widely acknowledged. They dominate the landscape of Cabo Verde, the southernmost and driest archipelago of Macaronesia, and are relevant natural resources for local populations, but a comprehensive evaluation of their distribution patterns is still lacking. In this study, we aim to evaluate the potential effects of climate change using the long-term data concerning grass distribution in Cabo Verde and the widely recognized climatic variability of this archipelago, which entails a huge irregularity in spatial and temporal rainfall. We identified two contrasting climatic periods (wet, from 1929 to 1968, and dry, from 1969 to 2007) and gathered all the information available from the bibliography, herbaria, and fieldwork concerning spontaneous grass species recorded in Cabo Verde during those two periods, which amounted to 107 taxa. This information was then used to disclose the patterns of grass diversity as related to climatic and topographic variables (altitude and windward vs. leeward aspects). Different altitudinal shifts in the distribution patterns of grass species assemblages and an assemblage specific to the wet period were revealed by comparing the two climatic periods. The role of exposure in delimiting the altitudinal distribution of the various assemblages was highlighted; the trade winds clearly determine the distribution of grass assemblages. We detected shifts in the distribution of grass assemblages according to the climatic periods (related to the macroclimate) and local topographic factors (associated with mesoclimates). Also, functional traits (i.e., annuals vs. perennials, C3 vs. C4 grasses, and tropical vs. temperate species) were found to vary between wet and dry periods, as well as with altitude and with slope aspect. Understanding species distributions and the role of the climatic variability of Cabo Verde is crucial to predicting how climate change will affect them and thus to support effective management and conservation actions.
Journal Article
Geographical isolation, habitat shifts and hybridisation in the diversification of the Macaronesian endemic genus Argyranthemum (Asteraceae)
• Inferring the processes responsible for the rich endemic diversity of oceanic island floras is important for our understanding of plant evolution and setting practical conservation priorities. This requires an accurate knowledge of phylogenetic relationships, which have often been difficult to resolve due to a lack of genetic variation.
• We employed genotyping-by-sequencing (GBS) to investigate how geographical isolation, habitat shifts, and hybridisation have contributed to the evolution of diversity observed in Argyranthemum Webb (Asteraceae), the largest genus of flowering plants endemic to the Macaronesian archipelagos.
• Species relationships were resolved, and biogeographical stochastic mapping identified intra-island speciation as the most frequent biogeographic process underlying diversification, contrary to the prevailing view in Argyranthemum and the Canary Islands. D-statistics revealed significant evidence of hybridisation between lineages co-occurring on the same island, however there was little support for the hypothesis that hybridisation may be responsible for the occurrence of nonmonophyletic multi-island endemic (MIE) species.
• Geographic isolation, habitat shifts and hybridisation have all contributed to the diversification of Argyranthemum, with intra-island speciation found to be more frequent than previously thought. Morphological convergence is also proposed to explain the occurrence of nonmonophyletic MIE species. This study reveals greater complexity in the evolutionary processes generating Macaronesian endemic diversity.
Journal Article
reconstruction of Palaeo-Macaronesia, with particular reference to the long-term biogeography of the Atlantic island laurel forests
Macaronesia is a biogeographical region comprising five Atlantic Oceanic archipelagos: the Azores, Madeira, Selvagen (Savage Islands), Canaries and Cape Verde. It has strong affinities with the Atlantic coast of the Iberian Peninsula and the north-western fringes of Africa. This paper re-evaluates the biogeographical history and relationships of Macaronesia in the light of geological evidence, which suggests that large and high islands may have been continuously available in the region for very much longer than is indicated by the maximum surface area of the oldest current island (27 Ma) - possibly for as long as 60 million years. We review this literature, attempting a sequential reconstruction of Palaeo-Macaronesia from 60 Ma to the present. We consider the implications of these geological dynamics for our understanding of the history of colonization of the present islands of Macaronesia. We also evaluate the role of these archipelagos as stepping stones and as both repositories of palaeo-endemic forms and crucibles of neo-endemic radiations of plant and animal groups. Our principal focus is on the laurel forest communities, long considered impoverished relicts of the Palaeotropical Tethyan flora. This account is therefore contextualized by reference to the long-term climatic and biogeographical history of Southern Europe and North Africa and by consideration of the implications of changes in land-sea configuration, climate and ocean circulation for Macaronesian biogeography. We go on to provide a synthesis of the more recent history of Macaronesian forests, which has involved a process of impoverishment of the native elements of the biota that has accelerated since human conquest of the islands. We comment briefly on these processes and on the contemporary status and varied conservation opportunities and threats facing these forests across the Macaronesian biogeographical region.
Journal Article
Biogeography and integrative taxonomy of Epipterygium (Mniaceae, Bryophyta)
A significant number of bryophyte species are thought to have transcontinental geographic ranges, oftenwithmultiple disjunct distribution areas. One of these cases is Epipterygiumtozeri (Mniaceae), with a Holarctic distribution and disjunct ranges inwestern North America, the Mediterranean, Japan and central Asia. Collections from different geographic regions were lumped into E. tozeri based on morphology, but a molecular confirmation was lacking so far. Here, we tested species concepts in the genus Epipterygium, with a special focus on the E. tozeri species complex, combining morphological and DNA sequence data for the nuclear ribosomal ITS region and two plastid loci (trnG intron, trnT-psbD spacer). In a second step, we reconstructed the historical biogeography of the genus. We found that Epipterygium most likely originated in Asia or North/Central America and that the alleged single widespread species E. tozeri with disjunct ranges is in fact a group of genetically and morphologically distinct taxa, including four overlooked species, for which we provide descriptions: E. atlanticum sp. nov., E. biauritum sp. nov., E. oreophilum sp. nov., and E. yunnanense sp. nov. The biogeographical history of these species is best explained by a step-wise parallel colonization of the Eurasian and American continents followed by in-situ speciation.
Journal Article
DNA barcoding of moon jellyfish (Cnidaria, Scyphozoa, Ulmaridae, Aurelia): two cryptic species from the Azores (ne atlantic, macaronesia), and evaluation of the non-íIndigenous species (NIS)
Moon jellies are some of the most popular, widely distributed, and best-studied marine jellyfish. By the end of the past century only two or three Aurelia species were recognized, but with the rise of DNA barcoding studies, around thirty Aurelia species are presently accepted. Most of the species are morphologically indistinguishable and have restricted biogeography. We reveal, with COI, 16S, and ITS1-5.8S sequence data, two (pseudo-)cryptic species of Aurelia, potentially endemic to the Azores ecoregion, herein provisionally classified as A. \"cf. pseudosolida\" and A. \"misteriosa\". These species are closely related to the Mediterranean lineages of A. pseudosolida and A. persea, respectively. In the Azores, the shape of the campanula and oral arms readily distinguishes the two species: the former with folded oral arms and globose campanula, and the latter with flattened campanula and thick and long oral arms. Previous reports of A. solida and A. aurita in the Azores should generally correspond to A. \"misteriosa\" and A. cf. pseudosolida, respectively. The phylogenetic (re-)examination of the available DNA barcodes of Aurelia only evidenced human-mediated dispersal for A. coerulea, A. relicta, and A. aurita. Aurelia solida cannot be yet considered NIS in the Mediterranean. More jellyfish DNA (meta)barcoding should reveal further cryptic diversity, biological invasions, and phylogeographic inferences.
Journal Article
Dietary Exposure to Toxic Metals (Cd, Pb and Hg) from Cereals Marketed in Madeira and the Azores
Cereals and cereal-based foods continue to be basic foods in all diets. Despite being known for their high nutritional value; they can also contain contaminants (hazards) such as toxic metals. This study assesses the Cd, Pb and Hg dietary exposure from cereals and derivatives marketed in Madeira and the Azores and characterizes the risks by evaluating the Cd and Hg intake contributions to the tolerable intakes and by estimating the Margin of Exposure (MOE) in the case of Pb. In Madeira, metals follow the descending order of Cd > Pb > Hg. Cd stands out as having the highest levels (0.307 mg Cd/kg in oats; 0.237 mg/kg in rye). High levels of Pb (0.347 mg/kg) were also detected in rye. Regarding total mercury, rice stands out (0.0013 mg/kg) followed by wheat (0.001 mg/kg). While all cereals and derivatives except maize consumed in Madeira exceed the maximum value of Cd allowed by the EU, 50.0% of the rye and 25.0% of the corn flour samples exceeded the European Pb limit. The daily consumption of 100 g of oats, rye flour and rye represent high contributions to the TWI of Cd (93.2 – 120%). The MOE values of Pb from the consumption of rye (100 g/day) are 1,294 (nephrotoxic effects) and 3,082 (cardiotoxic effects). In the Azores, corn flour (0.72 mg Pb/kg) stands out with 85.7% of the samples exceeding the maximum Pb EU limit and MOE values of 626 (nephrotoxic effects) and 1,490 (cardiotoxic effects). Regular daily consumption of corn flour makes a low (< 10%) contribution to the Cd TDI. In conclusion, the Pb exposure from the consumption of cereals and derivatives could have toxic effects such as nephrotoxicity or cardiotoxicity in adults. The results highlight the need to set up monitoring and surveillance programs for the safety of cereals and their derivatives in Madeira and the Azores in terms of lead and cadmium.
Journal Article
An updated checklist of Azorean arthropods (Arthropoda)
The Azores is a remote oceanic archipelago of nine islands which belongs to the Macaronesia biogeographical region hosting a unique biodiversity. The present Azorean landscape is strongly modified by the presence of man and only in small areas, where the soil or climate was too rough, have primitive conditions remained unchanged. Despite the fact that most of the Azorean native habitats are now lost, a large number of endemic species are still present and need urgent conservation. The present checklist of terrestrial and freshwater arthropods of the Azores Archipelago is based on all known published literature. The main goal of this work is to list, as rigorously as possible, all the known terrestrial and freshwater arthropods of the Azores. In this way, we are contributing to solve the ‘Linnaean’ shortfall, i.e. an incomplete taxonomic description of species-level diversity and the Wallacean Biodiversity Shortfall, the incomplete species distribution knowledge. The checklist includes new records of arthropods at island and archipelago levels that were published in the last twelve years. Compared to the last checklist of Azorean arthropods (Borges et al. 2010b), a total of 217 taxa (species and subspecies) are added. Currently, the total number of terrestrial and freshwater arthropod species and subspecies in the Azores is estimated to be 2420 taxa belonging to 14 classes, 53 orders, 440 families, 1556 genera, 2400 species and 149 individual subspecies. The most diverse orders of Azorean arthropods are: Coleoptera (585 taxa), Diptera (423 taxa), Hemiptera (338 taxa), Hymenoptera (163 taxa), Lepidoptera (159 taxa) and Araneae (133 taxa). A total of 276 endemic taxa are currently known (232 species and 44 subspecies), belonging to eight classes and 22 orders. São Miguel, Terceira and Pico are the islands with higher number of endemic species and subspecies. In the Azores, the number of native non-endemic taxa is 793 taxa, totalling 1069 indigenous taxa. Compared to the other nearest Macaronesian archipelagos (Madeira and Canaries), the Azorean arthropod fauna is characterised by a lower percentage of endemism (endemics/indigenous: 26% in Azores, 47% in Madeira Archipelago and 42% in Canary Islands) and a high proportion of exotic introduced taxa (39% in Azores, 19% in Madeira Archipelago and 8% in Canary Islands). Based on recent IUCN Red-listing of Azorean arthropods, a large fraction of the endemic taxa is under high threat.
Journal Article