Origins of the central Macaronesian psyllid lineages (Hemiptera; Psylloidea) with characterization of a new island radiation on endemic Convolvulus floridus (Convolvulaceae) in the Canary Islands.

A molecular survey of native and adventive psyllids in the central Macaronesian islands provides the first comprehensive phylogenetic assessment of the origins of the psyllid fauna of the Canary and Madeira archipelagos. We employ a maximum likelihood backbone constraint analysis to place the central Macaronesian taxa within the Psylloidea mitogenome phylogeny. The native psyllid fauna in these central Macaronesian islands results from an estimated 26 independent colonization events. Island host plants are predicted by host plants of continental relatives in nearly all cases and six plant genera have been colonized multiple times (Chamaecytisus, Convolvulus, Olea, Pistacia, Rhamnus, and Spartocytisus) from the continent. Post-colonization diversification varies from no further cladogenesis (18 events, represented by a single native taxon) to modest in situ diversification resulting in two to four native taxa and, surprisingly, given the diverse range of islands and habitats, only one substantial species radiation with more than four native species. Specificity to ancestral host plant genera or family is typically maintained during in situ diversification both within and among islands. Characterization of a recently discovered island radiation consisting of four species on Convolvulus floridus in the Canary Islands shows patterns and rates of diversification that reflect island topographic complexity and geological dynamism. Although modest in species diversity, this radiation is atypical in diversification on a single host plant species, but typical in the primary role of allopatry in the diversification process.

A review of the jumping plant-lice (Hemiptera: Psylloidea) of the Canary Islands, with descriptions of two new genera and sixteen new species.

The psyllid fauna of the Canary Islands is reviewed on the basis of recent field work on Tenerife, La Gomera, La Palma Gran Canaria and Lanzarote, as well as the examination of material deposited in several collections. Two new genera and 16 new species are described: Drepanoza Bastin, Burckhardt & Ouvrard gen. nov., Percyella Bastin, Burckhardt & Ouvrard gen. nov., Agonoscena atlantica Bastin, Burckhardt & Ouvrard sp. nov., A. sinuata Bastin, Burckhardt & Ouvrard sp. nov., Arytaina meridionalis Bastin, Burckhardt & Ouvrard sp. nov., Cacopsylla crenulatae Bastin, Burckhardt & Ouvrard sp. nov., C. falcicauda Bastin, Burckhardt & Ouvrard sp. nov., Diaphorina gonzalezi Bastin, Burckhardt & Ouvrard sp. nov., Drepanoza canariensis Bastin, Burckhardt & Ouvrard sp. nov., D. fruticulosi Bastin, Burckhardt & Ouvrard sp. nov., D. molinai Bastin, Burckhardt & Ouvrard sp. nov., Euphyllura confusa Bastin, Burckhardt & Ouvrard sp. nov., Percyella benahorita Bastin, Burckhardt & Ouvrard sp. nov., P. canari Bastin, Burckhardt & Ouvrard sp. nov., P. gomerita Bastin, Burckhardt & Ouvrard sp. nov., P. guanche Bastin, Burckhardt & Ouvrard sp. nov., Strophingia canariensis Bastin, Burckhardt & Ouvrard sp. nov. and S. paligera Bastin, Burckhardt & Ouvrard sp. nov. Arytaina insularis Loginova, 1976, stat. nov. is elevated from a subspecies of A. devia Loginova, 1976 to species rank. Five new combinations are proposed: Drepanoza fernandesi (Aguiar, 2001) comb. nov., D. lienhardi (Burckhardt, 1981) comb. nov., D. montanetana (Aguiar, 2001) comb. nov., D. pittospori (Aguiar, 2001) comb. nov. and Lauritrioza laurisilvae (Hodkinson, 1990) comb. nov., all transferred from Trioza. The hitherto unknown last-instar immatures are described for Megadicrania tecticeps Loginova, 1976, Cacopsylla atlantica (Loginova, 1976) and Lauritrioza laurisilvae (Hodkinson, 1990). We also report Agonoscena targionii (Lichtenstein, 1874) for the first time from the Canary Islands. Our review increases the number of known psyllid taxa from the Canary Islands to 24 genera and 62 species (one of which remains undescribed). Endemism, host plant relationships and biogeographic patterns are discussed. Keys for the identification of adults and immatures are provided, as well as information on host plants, distribution and predators.

Taxonomy and nomenclature of Abutilonalbidum (Malvaceae, Malvoideae), a cryptic Saharo-Canarian species recently rediscovered in Tenerife.

Abutilonalbidum, a cryptic Saharo-Canarian species, was thought to have been last collected in 1945 in Tenerife by E.R. Sventenius. In 2019, it was rediscovered in the same area. The characteristic features of the Canarian plants are discussed, especially in relation to the morphologically similar-looking and probably closely-related species Abutilonindicum and A.bidentatum. It is concluded that the plants from Tenerife and north-western Africa indeed represent a distinct species. The species is illustrated and a key for the identification of this and related species is presented.

How harmful are exotic plantations for soils and its microbiome? A case study in an arid island.

The plantation of exotic species has been a common practice in (semi-) arid areas worldwide aiming to restore highly degraded habitats. The effects of these plantations on plant cover or soil erosion have been widely studied, while little attention has been paid to the consequences on soil quality and belowground biological communities. This study evaluates the long-term (>60 years) effects of the exotic species Acacia cyclops and Pinus halepensis revegetation on soil properties, including microbiome, in an arid island. Soils under exotic plantation were compared to both degraded soils with a very low cover of native species and soils with well-preserved native plant communities. Seven scenarios were selected in a small area (~25 ha) with similar soil type but differing in the plant cover. Topsoils (0-15 cm) were analyzed for physical, chemical and biochemical properties, and amplicon sequencing of bacterial and fungal communities. Microbial diversity was similar among soils with exotic plants and native vegetation (Shannon's index = 5.26 and 5.34, respectively), while the most eroded soils exhibited significantly lower diversity levels (Shannon's index = 4.72). Bacterial and fungal communities' composition in degraded soils greatly differed from those in vegetated soils (Canberra index = 0.85 and 0.92, respectively) likely due to high soil sodicity, fine textures and compaction. Microbial communities' composition also differed in soils covered with exotic and native species, to a greater extent for fungi than for bacteria (Canberra index = 0.94 and 0.89, respectively), due to higher levels of nutrients, microbial biomass and activity in soils with native species. Results suggest that reforestation succeeded in avoiding further soil degradation but still leading to relevant changes in soil microbial community that may have negative effects on ecosystem stability. Information gained in this research could be useful for environmental agencies and decision makers about the controversial replacement of exotic plants in insular territories.

Reconstruction of the spatio-temporal diversification and ecological niche evolution of Helianthemum (Cistaceae) in the Canary Islands using genotyping-by-sequencing data.

BACKGROUND AND AIMS: Several biogeographical models have been proposed to explain the colonization and diversification patterns of Macaronesian lineages. In this study, we calculated the diversification rates and explored what model best explains the current distribution of the 15 species endemic to the Canary Islands belonging to Helianthemum sect. Helianthemum (Cistaceae). METHODS: We performed robust phylogenetic reconstructions based on genotyping-by-sequencing data and analysed the timing, biogeographical history and ecological niche conservatism of this endemic Canarian clade. KEY RESULTS: Our phylogenetic analyses provided strong support for the monophyly of this clade, and retrieved five lineages not currently restricted to a single island. The pristine colonization event took place in the Pleistocene (~1.82 Ma) via dispersal to Tenerife by a Mediterranean ancestor. CONCLUSIONS: The rapid and abundant diversification (0.75-1.85 species per million years) undergone by this Canarian clade seems the result of complex inter-island dispersal events followed by allopatric speciation driven mostly by niche conservatism, i.e. inter-island dispersal towards niches featuring similar environmental conditions. Nevertheless, significant instances of ecological niche shifts have also been observed in some lineages, making an important contribution to the overall diversification history of this clade.

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.

Maximize Resolution or Minimize Error? Using Genotyping-By-Sequencing to Investigate the Recent Diversification of Helianthemum (Cistaceae).

A robust phylogenetic framework, in terms of extensive geographical and taxonomic sampling, well-resolved species relationships and high certainty of tree topologies and branch length estimations, is critical in the study of macroevolutionary patterns. Whereas Sanger sequencing-based methods usually recover insufficient phylogenetic signal, especially in recently diversified lineages, reduced-representation sequencing methods tend to provide well-supported phylogenetic relationships, but usually entail remarkable bioinformatic challenges due to the inherent trade-off between the number of SNPs and the magnitude of associated error rates. The genus Helianthemum (Cistaceae) is a species-rich and taxonomically complex Palearctic group of plants that diversified mainly since the Upper Miocene. It is a challenging case study since previous attempts using Sanger sequencing were unable to resolve the intrageneric phylogenetic relationships. Aiming to obtain a robust phylogenetic reconstruction based on genotyping-by-sequencing (GBS), we established a rigorous methodological workflow in which we i) explored how variable settings during dataset assembly have an impact on error rates and on the degree of resolution under concatenation and coalescent approaches, ii) assessed the effect of two extreme parameter configurations (minimizing error rates vs. maximizing phylogenetic resolution) on tree topology and branch lengths, and iii) evaluated the effects of these two configurations on estimates of divergence times and diversification rates. Our analyses produced highly supported topologically congruent phylogenetic trees for both configurations. However, minimizing error rates did produce more reliable branch lengths, critically affecting the accuracy of downstream analyses (i.e. divergence times and diversification rates). In addition to recommending a revision of intrageneric systematics, our results enabled us to identify three highly diversified lineages in Helianthemum in contrasting geographical areas and ecological conditions, which started radiating in the Upper Miocene.

Are there any widespread endemic flowering plant species in Macaronesia? Phylogeography of Ranunculus cortusifolius.

PREMISE OF THE STUDY: Oceanic island endemics typically exhibit very restricted distributions. In Macaronesia, only one endemic angiosperm species, Ranunculus cortusifolius, has a distribution spanning the archipelagos of the Azores, Madeira, and Canaries. Earlier work suggested possible differences between archipelagos and the multiple origins of the species. This paper tests the hypothesis that R. cortusifolius is a single widespread Macaronesian endemic species with a single origin. METHODS: Chloroplast (matK-trnK, psbJ-petA) and ITS sequences were generated from across the distribution of R. cortusifolius. Relationships were investigated using Bayesian inference and divergence times estimated using BEAST. Infraspecific variation was investigated using statistical parsimony. The general mixed Yule-coalescent model (GMYC) was further used to identify putative species boundaries based on maternally inherited plastid data. KEY RESULTS: The hypothesis of multiple independent origins of R. cortusifolius is rejected. Divergence of the R. cortusifolius lineage from a western Mediterranean sister group in the late Miocene is inferred. Distinct genotypes were resolved within R. cortusifolius that are endemic to the Azores, Madeira, and the Canaries. Four to five putative species were delimited by different versions of the GMYC model. CONCLUSION: Ranunculus cortusifolius is the result of a single colonization of Macaronesia. The large distances between archipelagos have been effective barriers to dispersal, promoting allopatric diversification at the molecular level with diversification also evident within the Canaries. Isolation has not been accompanied by marked morphological diversification, which may be explained by the typical association of R. cortusifolius with stable and climatically buffered laurel forest communities.

Molecular taxonomic identification in the absence of a 'barcoding gap': a test with the endemic flora of the Canarian oceanic hotspot.

We use a comprehensive subset of Canarian angiosperms corresponding to 23 families, 35 genera and 60 Canarian endemic taxa to test whether this flora is suitable to taxonomic identification with the two proposed plant DNA barcode sequences and whether these sequences may reveal the existence of cryptic species overlooked by morphology. The rate of discrimination success between the insular congeneric samples using the rbcL+matK combination and a 'character-based' approach (where we use only the combination of nucleotide positions in an alignment that allows unambiguous species identification) is higher (82.29%) than that obtained with the 'distance-based' approach (80.20%) used by the CBOL Plant Working Group in 2009 and also when compared with tests conducted in other floras. This suggests that the molecular identification of the Canarian endemic flora can be achieved as successfully as in other floras where the incidence of radiation is not as relevant. The facts that (i) a distance-based criterion was unable to discriminate between congeneric and conspecific comparisons and (ii) only the character-based discrimination criterion resolved cases that the distance-based criterion did not, further support the use of a character discrimination approach for a more efficient DNA barcoding of floras from oceanic islands like the Canaries. Thus, a barcoding gap seems not to be necessary for the correct molecular characterization of the Canarian flora. DNA barcodes also suggest the possible existence of cryptic taxa to be further investigated by morphology and that the current taxonomic status of some of the taxa analysed may need revision.

Allopatric diversification, multiple habitat shifts, and hybridization in the evolution of Pericallis (Asteraceae), a Macaronesian endemic genus.

PREMISE OF THE STUDY: Geographic isolation, habitat shifts, and hybridization have contributed to the diversification of oceanic island floras. We investigated the contribution of these processes to the diversification of Pericallis, a genus endemic to Macaronesia. METHODS: Data from the chloroplast psaI-accD and trnV-ndhC regions and the nuclear ribosomal internal transcribed spacer region (ITS) were sampled for multiple accessions of all taxa and used to establish phylogenetic hypotheses. Habitat preferences were optimized to investigate habitat shifts, and divergence times were estimated. Species nonmonophyly was investigated using Bayes factors. KEY RESULTS: Much of the diversification in Pericallis has occurred recently, within the past 1.7 Ma. Three habitat shifts have occurred in the evolution of the genus. However, geographic isolation has played a greater role in its diversification. Novel allopatric patterns were revealed within some species, highlighting the significance of geographic isolation in the evolution of Pericallis. One species (P. appendiculata) that resolved as monophyletic in the ITS analysis was polyphyletic in the chloroplast analysis. Bayes factors provide strong support for the nonmonophyly of P. appendiculata haplotypes, and their phylogenetic placement suggests that ancient hybridization is responsible for the haplotype diversity observed. CONCLUSIONS: Multiple markers and extensive sampling provided new insights into the evolution of Pericallis. In contrast to previous studies, our results reveal a more significant role for allopatry than habitat shifts and new evidence for ancient hybridization in the evolution of Pericallis. Our study highlights the power of broad taxon sampling for unraveling diversity patterns and processes within oceanic island radiations.