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[Aim] One of the main goals of functional biogeography is to examine distribution patterns of trait diversity, and islands provide excellent study cases for this emerging field. We tested the hypothesis that multiple dispersals from a common mainland pool would promote functional similarity among island systems when environmental conditions are similar, but also novel phenotypic traits related to colonization history and exploitation of new habitats., [Location] Mediterranean Basin and Macaronesian islands., [Taxon] Wolfbane (Periploca laevigata), [Methods] We used the well‐known biogeographical history of a woody plant complex (P. laevigata s.l.) to examine trait variation and how it relates to climatic conditions of mainland and subtropical island settings. In a common garden experiment, we measured a suite of leaf physiological and anatomical traits tightly related to plant performance in 320 seedlings representing 21 populations of five sublineages—the oldest (2.6 my) island colonization (western Canary Islands) as a reference, three sublineages stemming from independent events of island colonization in the last 0.5 my from NW Africa (Cape Verde, Fuerteventura, Lanzarote) and their widespread Mediterranean mainland counterpart., [Results] We observed strong phenotypic divergence between island and mainland sublineages linked to contrasting climatic conditions. Mediterranean mainland populations displayed a very specialized leaf phenotype characteristic of arid plants (i.e. small leaves, amphistomaty, isobilateral mesophyll, high photosynthetic rates). In turn, low seasonality on islands was linked to the recurrent expression of a phenotype characterized by larger leaves and lower photosynthetic rates. Our analyses showed that the high investment in secondary compounds (i.e. tannins) on islands decouples photosynthesis from growth rates. Despite this pattern of parallel differentiation, each island sublineage displayed a distinctive phenotype, with some traits related to colonization time, which resulted in a mosaic of functional variation across island systems., [Main conclusions] Our data suggest that the studied subtropical islands promote expression of traits specific to certain sublineages and other common traits that are no longer adaptive in the original mainland pool due to Pleistocene climatic shifts. These findings ultimately extend the role of islands as biodiversity refugia and hotspots of plant functional diversity., This research was funded by the European Regional Development Fund (Intereg projects CAVEGEN and ENCLAVES). C. G‐V was financially supported by a ‘Vicenç Mut' postdoctoral fellowship (Conselleria d'Innovació, Recerca i Turisme, Govern de les Illes Balears and the European Social Fund).

Subtropical islands are often viewed as refuges where Quaternary climatic shifts driving global episodes of extinction were buffered. Island biodiversity, however, may have been impacted by climatic fluctuations at local scales, particularly in spatially heterogeneous island systems. In this study, we generated a conceptual framework for predicting the potential impact of Pleistocene extinctions on the biogeographical pattern of the Canarian spermatophyte flora, with a focus on the easternmost Canarian islands (ECI). Then, we performed an exhaustive bibliographic revision (270 studies) to examine whether taxonomic, phylogenetic and phylogeographical data support our predictions. Although molecular information is limited for many lineages, the available data suggest that the majority of extant ECI plant taxa may be the result of relatively recent (, C. G.V. was funded by project SV‐17‐GIJON‐BOTANICO. J.P. was funded by the MINECO through the Ramón y Cajal Program (RYC‐2016‐20506), and Marie Sklodowska‐Curie COFUND, Researchers' Night and Individual Fellowships Global (MSCA grant agreement No 747238, 'UNISLAND').

[Aim] Plants on islands are often subjected to lower levels of herbivory relative to those found at mainland sites. As a consequence, island plants are predicted to exhibit lower levels of physical or chemical defences, which renders them more susceptible to introduced herbivores. Yet, instances of high pressure by superabundant herbivores native to islands have been reported in many insular systems, which presumably would result in heightened plant defences. To date, no quantitative review has been conducted to determine how common these contrasting patterns are and their implications for the evolution of plant-herbivore interactions., [Location] Islands worldwide., [Taxon] Plants, insects, molluscs, mammals., [Methods] We conducted a meta-analysis of insularity effects on herbivory and plant defences by including studies that involved island-mainland comparisons of the same plant species in both environments (90% of cases), or insular endemics versus mainland congeners (10% of cases). We tested for differences between mammalian and invertebrate (mollusc or insect) herbivory as well as between plant chemical and physical defences by specifying comparisons based on the type of herbivore (vertebrate or invertebrate) or plant trait included in the study., [Results] Mammalian herbivory was significantly higher on islands than on mainlands. In contrast, no significant effect was observed on invertebrate herbivory. In addition, we found no significant difference in either plant physical or chemical defences between insular and mainland plants, though physical defences tended to be higher for plants on islands., [Main conclusions] All analysed mammal studies focused on species introduced to islands, suggesting greater susceptibility of insular plants to exotic mammals, whereas the lack of effects in the case of invertebrate herbivory suggests no difference in susceptibility to molluscs or insects between insular and mainland plants. Interestingly, plant trait patterns suggest a trend for increased physical defences by insular plants, possibly due to heightened pressure by exotic mammalian herbivores on islands, whereas chemical defences appear uncorrelated to differences in herbivory. These findings call for further experimental and observational studies measuring defences and herbivory for multiple sympatric plant species occurring at both mainland and island sites within a system, or comparing insular endemics to congeneric mainland species., This research was financially supported by a grant from the Regional Government of Galicia (IN607D 2016/001), a grant from the Spanish Ministry of Science, Innovation and Universities (RTI2018-099322-B-100) and the Ramón y Cajal Research Programme (RYC-2013-13230) to XM. CG-V is supported by a postdoctoral Vicenç Mut fellowship (Govern de les Illes Balears, Conselleria d'Innovació, Recerca i Turisme and the European Social Fund).

Accurate inference in phylogeography requires appropriate sampling strategies. Complex questions demand a large sample size at both the population and genetic levels to obtain precise reconstructions. This is the case of the phylogeographic history of Cistus monspeliensis, a plant that displays low plastid (cpDNA) diversity in the Mediterranean Basin but high diversity in the Canary Islands. Here, we aimed to identify Mediterranean refugial areas and to accurately quantify inter-island colonization events in the Canaries. Using a previous study as starting point, we increased sample size in two ways: (i) additional sampling of plastid genetic markers (from 1041 to 1899 bp); and (ii) additional sampling of populations (from 47 to 69) in long-term persistence areas suggested by species distribution modeling (SDM). The synergy between SDM and extended population sampling helped find higher genetic diversity. Our deeper phylogeographic sampling of C. monspeliensis revealed the following: (i) potential refugia in long-term persistence areas with high cpDNA diversity in western Europe and the Canary Islands; and (ii) a significant increase (from 7 to 12) in the number of inferred inter-island colonization events across the archipelago. Our results stress the usefulness of SDM to identify the genetic signature associated with potential refugial areas. We herein propose a field sampling approach based on SDM that, in combination with a larger cpDNA sampling, can help answer a wide array of phylogeographic questions, such as the location of Quaternary refugia and number of colonizations across archipelagos.

The genus Dracaena in Macaronesia comprises two threatened species of arborescent monocots that are often associated with one of the most intriguing biogeographic disjunctions: the Rand Flora pattern. Molecular information is, however, largely missing for the Macaronesian Dracaena taxa (“MDT”, hereafter), and the biogeographic or population genetic patterns of this lineage have not yet been thoroughly assessed. To fill this gap, we generated plastid DNA sequence data of 14 Dracaena populations representing the entire natural distribution of MDT (including mainland Morocco and all recognized subspecies), 9 additional populations of subspontaneous origin, and a set of related species of the genus. We performed phylogenetic, biogeographic, and population genetic analyses at different spatial scales and conducted a comparative review on plant haplotype diversity in Macaronesian plants. The results of our phylogenetic analyses indicated the monophyly of the MDT and an origin separate from a clade of geographically distant species that so far were postulated as their closest living relatives (D. cinnabari, D. ombet, D. schizantha, D. serrulata). The results of our phylogeographic analyses indicated that diversification within D. draco occurred throughout the Pleistocene and that wild peripheral populations (Madeira, mainland Morocco) may have a recent origin from Canarian source populations. Recent dispersals, coupled with remarkably low levels of haplotype diversity, probably account for the weak phylogeographic signal observed across wild populations. However, our results suggested that human‐assisted expansion of Dracaena inflates the extant phylogeographic signal by non‐random translocation of a specific subset of haplotypes. Our study demonstrates that many of the previous biogeographic scenarios on MDT are not supported by molecular data. Instead, our results highlight (i) the impact that human activity may have on the phylogeographic pattern of island plants, and (ii) the need of a deeper taxonomic sampling in future investigations on MDT and close relatives., This research was funded by the European Regional Development Fund (Intereg projects CAVEGEN and ENCLAVES). JP was funded by the Marie Sklodowska‐Curie COFUND, Researchers’ Night and Individual Fellowships Global (MSCA grant agreement No 747238, “UNISLAND”’), and the Ramón y Cajal program (RYC‐2016‐20506). CG‐V was financially supported by a “Vicenç Mut” postdoctoral fellowship (Conselleria d'Innovació, Recerca i Turisme, Govern de les Illes Balears and the European Social Fund).

We use the molecular data available for the plant genera currently distributed in the Canaries to propose a hypothetical framework that updates the Surfing Syngameon Hypothesis (SSH) to explain the origins of the high genetic diversity of this flora in connection with the General Dynamic Model of Island Ontogeny and the available knowledge on the geology of the islands. Based on our review, we propose that (i) secondary contact and subsequent gene flow among genotypes previously confined to the mainland or to other insular regions generated syngameons across the archipelago, whose temporal extent and distribution were influenced by the ontogenetic stages of each island; and (ii) the resulting high levels of genetic variation were critical to the successful colonization of many other regions in the Canaries, especially those where more complex geographical and ecological features promoted gene flow cessation, drift, and/or eventual selection processes leading to species radiations. Under the updated SSH, the colonizing contingents of the islands would not necessarily have low genetic variation, because multiple colonizations followed by secondary contact among different genotypes would enhance genetic diversity through introgressive hybridization. As with other theories, the SSH contends that the monophyly detected in a large proportion of its lineages is artifactual. However, the SSH further differs from past hypotheses in (i) suggesting a spatio-temporal sequence of events giving rise to syngameons in different islands and island regions, (ii) showing that the ontogenetic mismatch among the Canarian islands should be largely responsible for similar levels of population genetic diversity in many congeneric endemics distributed in several islands, (iii) hypothesizing a substantial role for both allopolyploid and homoploid hybridization in the origins of the Canarian flora, and (iv) positing that, in many cases, the admixture processes in the syngameons formed at different stages of the islands’ ontogeny made possible the rapid colonization of multiple environments, and the generation of the current floristic diversity.

Strong geographical isolation within the distribution of a species may result in differentiated lineages exhibiting conspicuous phenotypic differences. In the present paper, we investigate whether plastid and phenotypic variation is geographically structured within the Olea europaea complex in Macaronesia, which comprises three subspecies separated by oceanic barriers: maroccana (south-west Morocco), guanchica (Canary Islands) and cerasiformis (Madeira archipelago). Plastid variation showed a significant pattern of geographical structure (NST > GST = 0.56), because of the lack of shared haplotypes among subspecies and the presence of a single and private haplotype in the eastern Canary Islands. Such a clear molecular structure, however, was not reflected in a congruent pattern of phenotypic differentiation among taxa in leaf morpho-functional traits. Despite the substantial genetic differentiation observed between the subspecies from Madeira and the Canary Islands, they displayed both higher leaf size (leaf area) and specific leaf area (leaf surface area-to-mass ratio) than their continental counterparts, probably as a result of oceanic conditions in subtropical environments. Unlike most of the plant groups previously studied in the Macaronesian region, the lineages of Olea illustrate how low phenotypic differentiation can be also related to a clear molecular differentiation in oceanic island enclaves. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162, 54–63.

Geographical isolation and polyploidization are central concepts in plant evolution. The hierarchical organization of archipelagos in this study provides a framework for testing the evolutionary consequences for polyploid taxa and populations occurring in isolation. Using amplified fragment length polymorphism and simple sequence repeat markers, we determined the genetic diversity and differentiation patterns at three levels of geographical isolation in Olea europaea: mainland-archipelagos, islands within an archipelago, and populations within an island. At the subspecies scale, the hexaploid ssp. maroccana (southwest Morocco) exhibited higher genetic diversity than the insular counterparts. In contrast, the tetraploid ssp. cerasiformis (Madeira) displayed values similar to those obtained for the diploid ssp. guanchica (Canary Islands). Geographical isolation was associated with a high genetic differentiation at this scale. In the Canarian archipelago, the stepping-stone model of differentiation suggested in a previous study was partially supported. Within the western lineage, an east-to-west differentiation pattern was confirmed. Conversely, the easternmost populations were more related to the mainland ssp. europaea than to the western guanchica lineage. Genetic diversity across the Canarian archipelago was significantly correlated with the date of the last volcanic activity in the area/island where each population occurs. At the island scale, this pattern was not confirmed in older islands (Tenerife and Madeira), where populations were genetically homogeneous. In contrast, founder effects resulted in low genetic diversity and marked genetic differentiation among populations of the youngest island, La Palma.

Interspecific introgression is a well-known phenomenon in oaks whose ecological and evolutionary consequences, although relevant, are still unclear. We investigated molecular variation and any evidence for hybridisation in nine natural populations of kermes oak ( Quercus coccifera L.) from the Iberian Peninsula. Additive patterns in 59 nrDNA ITS sequences revealed that hybrid individuals showing intermediate genotypes between kermes and holm ( Q. ilex L.) oaks are very frequent, although intermediate morphotypes are uncommon. Bayesian analysis of ISSR fingerprinting patterns indicated extensive gene-flow among Q. coccifera populations and neighbouring Quercus taxa. Introgression appeared to be an active mechanism in the pair Q. coccifera – Q. ilex , but no conclusive evidence supporting hybridisation between Q. coccifera and other co-occurring Quercus species was found. The role of canalisation in the maintenance of stable morphological characters in the face of extensive introgression is discussed.

†Background and Aims Quercus coccifera, as a long-lived sprouter, responds plastically to environmental variation. In this study, the role of foliar plasticity as a mechanism of habitat selection and modification within the canopy and across contrasted habitats was characterized. An examination was made of the differential contribution of inner and outer canopy layers to the crown plasticity expressed in the field by adult individuals and its dependence on environmental and genetic factors. †Methods Within-crown variation in eight foliar traits was examined in nine populations dominated by Q. coccifera. The difference between mean trait values at the inner and outer canopy layers was used as a proxy for crown plasticity to light. Correlations between geographic distances, environmental differences (climatic and edaphic) and phenotypic divergence (means and plasticities) were assessed by partial Mantel tests. A subset of field measurements was compared with data from a previous common garden experiment. †Key Results Phenotypic adjustment of sun leaves contributed significantly to the field variation in crown plasticity. Plasticity in leaf angle, lobation, xanthophyll cycle pigments and b-carotene content was expressed in sun and shade leaves concurrently and in opposite directions. Phenotypic plasticity was more strongly correlated with environmental variation than mean trait values. Populations of taller plants with larger, thinner (higher specific leaf area) and less spiny leaves exhibited greater plasticity. In these populations, the midday light environment was more uniform at the inner than at the outer canopy layers. Field and common garden data ranked populations in the same order of plasticity. †Conclusions The expression of leaf plasticity resulted in a phenotypic differentiation that suggests a mechanism of habitat selection through division of labour across canopy layers. Signs of plasticity-mediated habitat modification were found only in the most plastic populations. Intracanopy plasticity was sensitive to environmental variation but also exhibited a strong genetic component.

Understanding how genetic diversity is structured on oceanic island taxa requires the integration of physical, biological and anthropomorphic factors. Founder effects coupled with limited dispersal over sea barriers typically result in low levels of genetic variation in island populations. In widespread species, restriction in gene flow across large areas leads to patterns of isolation by distance (IBD), but recent population-based studies indicate that genetic structure on islands can be complex even at local scales. Here, we investigated the patterns of genetic variation in a widespread island palm (Phoenix canariensis) displaying reproductive syndromes associated with extensive dispersal (wind pollination and zoochory). Genetic variation was assessed at eight nuclear microsatellite loci in 330 individuals of 15 Canarian populations. Our results showed that levels of within-population genetic diversity in P. canariensis depend on the island considered, with a strong decreasing pattern from the easternmost and oldest island to the westernmost and youngest islands. A Mantel test supported a stepping-stone model of differentiation across the archipelago that fits the sequence of island emergence, and results from ABC and clustering analyses also corresponded with this progression rule. In addition, our analyses were congruent with the idea that the only large population found on Lanzarote has an anthropogenic origin. Despite the high dispersal potential of P. canariensis, our analyses suggest that the geographical configuration of the Canary Islands and a relatively recent pattern of differentiation across islands appear to have had a primary influence on the genetic structure of this island taxon.

Evergreen oaks are an emblematic element of the Mediterranean vegetation and have a leaf phenotype that seems to have remained unchanged since the Miocene. We hypothesise that variation of the sclerophyll phenotype among Iberian populations of Quercus coccifera is partly due to an ulterior process of ecotypic differentiation. We analysed the genetic structure of nine Iberian populations using ISSR fingerprints, and their leaf phenotypes using mean and intracanopy plasticity values of eight morphological (leaf angle, area, spinescence, lobation and specific area) and biochemical traits (VAZ pool, chlorophyll and beta-carotene content). Climate and soil were also characterised at the population sites. Significant genetic and phenotypic differences were found among populations and between NE Iberia and the rest of the populations of the peninsula. Mean phenotypes showed a strong and independent correlation with both genetic and geographic distances. Northeastern plants were smaller, less plastic, with smaller, spinier and thicker leaves, a phenotype consistent with the stressful conditions that prevailed in the steppe environments of the refugia within this geographic area during glaciations. These genetic, phenotypic, geographic and environmental patterns are consistent with previously reported palaeoecological and common evidence. Such consistency leads us to conclude that there has been a Quaternary divergence within the sclerophyllous syndrome that was at least partially driven by ecological factors.