Genetically depauperate in the continent but rich in oceanic islands: Cistus monspeliensis (Cistaceae) in the Canary Islands

Background

Population genetic theory holds that oceanic island populations are expected to have lower levels of genetic variation than their mainland counterparts, due to founder effect after island colonization from the continent. Cistus monspeliensis (Cistaceae) is distributed in both the Canary Islands and the Mediterranean region. Numerous phylogenetic results obtained in the last years allow performing further phylogeographic analyses in Cistus.

Methodology/Principal Findings

We analyzed sequences from multiple plastid DNA regions in 47 populations of Cistus monspeliensis from the Canary Islands (21 populations) and the Mediterranean basin (26 populations). The time-calibrated phylogeny and phylogeographic analyses yielded the following results: (1) a single, ancestral haplotype is distributed across the Mediterranean, whereas 10 haplotypes in the Canary Islands; (2) four haplotype lineages are present in the Canarian Islands; (3) multiple colonization events across the archipelago are inferred; (4) the earliest split of intraspecific lineages occurred in the Early to Middle Pleistocene (<930,000 years BP).

Conclusions/Significance

The contrasting pattern of cpDNA variation is best explained by genetic bottlenecks in the Mediterranean during Quaternary glaciations, while the Canarian archipelago acted as a refugium of high levels of genetic diversity. Active colonization across the Canarian islands is supported not only by the distribution of C. monspeliensis in five of the seven islands, but also by our phylogeographic reconstruction in which unrelated haplotypes are present on the same island. Widespread distribution of thermophilous habitats on every island, as those found throughout the Mediterranean, has likely been responsible for the successful colonization of C. monspeliensis, despite the absence of a long-distance dispersal mechanism. This is the first example of a plant species with higher genetic variation among oceanic island populations than among those of the continent.     

Interpreting colonization of the Calathus (Coleoptera: Carabidae) on the Canary Islands and Madeira through the application of the parametric bootstrap

The Canary Islands have proven to be an interesting archipelago for the phylogeographic study of colonization and diversification with a number of recent studies reporting evolutionary patterns and processes across a diversity of floral and faunal groups. The Canary Islands differ from the Hawaiian and Galapagos Islands by their close proximity to a continental land mass, being 110 km from the northwestern coast of Africa. This close proximity to a continent obviously increases the potential for colonization, and it can be expected that at the level of the genus some groups will be the result of more than one colonization. In this study we investigate the phylogeography of a group of carabid beetles from the genus Calathus on the Canary Islands and Madeira, located 450 km to the north of the Canaries and 650 km from the continent. The Calathus are well represented on these islands with a total of 29 species, and on the continent there are many more. Mitochondrial cytochrome oxidase I and II sequence data has been used to identify the phylogenetic relationships among the island species and a selection of continental species. Specific hypotheses of monophyly for the island fauna are tested with parametric bootstrap analysis. Data suggest that the Canary Islands have been colonized three times and Madeira twice. Four of these colonizations are of continental origin, but it is possible that one Madeiran clade may be monophyletic with a Canarian clade. The Calathus faunas of Tenerife and Madeira are recent in origin, similar to patterns previously reported for La Gomera, El Hierro, and Gran Canaria.     

Phylogeography and molecular phylogeny of Macaronesian island Tarphius (Coleoptera: Zopheridae): why are there so few species in the Azores?

Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra‐island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra‐island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra‐island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species‐rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature.     

Molecular evidence for adaptive radiation of Micromeria Benth. (Lamiaceae) on the Canary Islands as inferred from chloroplast and nuclear DNA sequences and ISSR fingerprint data

The Canary Islands have been a focus for phylogeographic studies on the colonization and diversification of endemic angiosperm taxa. Based on phylogeographic patterns, both inter island colonization and adaptive radiation seem to be the driving forces for speciation in most taxa. Here, we investigated the diversification of Micromeria on the Canary Islands and Madeira at the inter- and infraspecific level using inter simple sequence repeat PCR (ISSR), the trnK-Intron and the trnT-trnL-spacer of the cpDNA and a low copy nuclear gene. The genus Micromeria (Lamiaceae, Mentheae) includes 16 species and 13 subspecies in Macaronesia. Most taxa are restricted endemics, or grow in similar ecological conditions on two islands. An exception is M. varia, a widespread species inhabits the lowland scrub on each island of the archipelago and could represent an ancestral taxon from which radiation started on the different islands. Our analyses support a split between the "eastern" islands Fuerteventura, Lanzarote and Gran Canaria and the "western" islands Tenerife, La Palma and El Hierro. The colonization of Madeira started from the western Islands, probably from Tenerife as indicated by the sequence data. We identified two lineages of Micromeria on Gomera but all other islands appear to be colonized by a single lineage, supporting adaptive radiation as the major evolutionary force for the diversification of Micromeria. We also discuss the possible role of gene flow between lineages of different Micromeria species on one island after multiple colonizations.     

Origin and evolution of the endemic genera of Gonosperminae (Asteraceae: Anthemideae) from the Canary Islands: evidence from nucleotide sequences of the internal transcribed spacers of the nuclear ribosomal DNA

The Gonosperminae (Asteraceae) are composed of three genera endemic to the Canary Islands (GONOSPERMUM: Less., and LUGOA: DC.) and southern Africa (INULANTHERA: K?llersj?), and they are considered an example of a floristic link between these two regions. Phylogenetic analyses of ITS sequences reveal that the Canarian genera are not sister to INULANTHERA: and do not support the monophyly of the Gonosperminae. These results, coupled with previous phylogenetic studies of other groups, suggest that many of the putative biogeographic links between Macaronesia and southeast Africa need to be evaluated by rigorous phylogenetic analyses. INULANTHERA: forms part of the basal southern African radiation of the Anthemideae, and therefore it is closely related to other taxa from this region. Maximum likelihood and weighted parsimony analyses support a monophyletic group in the Canary Islands, that includes LUGOA:, Gonospermum, and three TANACETUM: species endemic to the island of Gran Canaria. Bootstrap support for the monophyly of this Canarian group is weak, and it collapses in the strict consensus tree based on unweighted parsimony. LUGOA: is nested within Gonospermum, and both interisland colonization among the western islands of La Gomera, El Hierro, La Palma and Tenerife, and radiation on the central island of Gran Canaria have been the major patterns of species diversification for these Canarian endemics.     

Evolution and diversification of the forest and hypogean ground-beetle genus Trechus in the Canary Islands

The beetle genus Trechus (Carabidae) is represented in the Macaronesian Islands by 43 endemic species. The Canary Islands have 16 endemic species, with two adapted to hypogean life. Phylogenetic relationships among 177 individuals of 38 Canarian, Madeiran, Azorean and continental Trechus species were examined using mitochondrial DNA and nuclear internal transcribed spacer 2 (ITS2) sequence data. Results show two main lineages in the Canaries: one comprising two sister groups with species from the laurel forest of La Gomera and Tenerife, and the other containing the single species from Gran Canaria and a species complex in the four western islands including two troglobites. Calibrations were applied to a linearized tree using a relaxed molecular clock method to estimate the major evolutionary divergence times of the Canarian Trechus species. Although the species assemblage in this archipelago is relatively ancient (7-8 million years), much of the species diversity is recent. Transition to the hypogean environment is more consistent with the "adaptive shift" rather than with the competing "climatic relict" hypothesis.     

Diversification of sympatric Sapromyza (Diptera: Lauxaniidae) from Madeira: six morphological species but only four mtDNA lineages

A series of recent studies on speciation of insects within the Canary Islands have indicated considerable within-island diversification, similar to that described in the Hawaiian islands. Little work has yet been carried out on the neighboring Madeiran archipelago, which is also volcanic. This study examines relationships among all known Lauxaniid flies of the genus Sapromyza from Madeira (including six newly described morphological species) based on mitochondrial gene trees constructed from cytochrome c oxidase (subunit I) and 16S rRNA partial sequences. Phylogenies based on maximum likelihood distances, a Bayesian method based on Markov chain Monte Carlo sampling from the posterior probability distribution, and maximum parsimony show that eight of the nine Madeiran species comprise a single monophyletic group. This clade is also split into two subclades representing black- and yellow/orange-bodied forms. The latter mtDNA clade corresponds to only two species (Sapromyza imitans and Sapromyza indigena) which are not reciprocally monophyletic. Monophyly is strongly supported within four of the six black-bodied species but not for the species pair (Sapromyza inconspicua, Sapromyza laurisilvae). We discuss the double occurrence (at least) of introgressive hybridization/incomplete lineage sorting within this group and suggest that recent speciation is the most likely explanation. The remaining species on the island, Sapromyza madeirensis, is very divergent from the aforementioned group, occupying a more basal position in the tree than the other Atlantic island and continental Sapromyza that were included in the analysis. At least two speciation events for Madeiran Sapromyza appear to correspond to quite ancient periods relative to the age of the island, while others are more recent. This suggests that a combination of island colonization and within-island sympatric and/or vicariance-mediated speciation may explain the observed diversity.     

Biogeographical and evolutionary patterns in the Macaronesian shield-backed katydid genus Calliphona Krauss, 1892 (Orthoptera: Tettigoniidae) and allies as inferred from phylogenetic analyses of multiple mitochondrial genes

Abstract.  We investigated the phylogenetic patterns, evolutionary processes, and their taxonomic implications, of two closely related shield-backed katydid genera endemic to the Macaronesian archipelagos: the monotypic Psalmatophanes Chopard, 1938 endemic to Madeira and Calliphona Krauss, 1892, which includes three species restricted to the Canary Islands. Two main hypotheses have been proposed to explain the origin and colonization pathways of these two genera: a single origin with subsequent sequential colonization of the islands, or three independent colonization waves from continental Africa. We used DNA sequence information from the mitochondrial genes cox1, tRNAleucine, rrnL and nad1 to infer phylogenetic relationships among Psalmatophanes and Calliphona species. Our results provide support for the independent colonization of Madeira and the Canary Islands, and suggest that Psalmatophanes is actually more closely related to the continental genus Tettigonia than to the Canarian representatives. Deep genetic divergence among Canarian species provides further support for the assignment of the Canarian species into two subgenera. Tree topology along with Bayesian-based estimates of lineage age suggest a pattern of colonization from Tenerife to La Palma, and from Tenerife to Gran Canaria with subsequent dispersal to La Gomera. We report the first collection of a Calliphona specimen in the island of El Hierro, which molecular data suggest is a recent immigrant from La Gomera. We hypothesize that the patterns of distribution and genetic divergence exhibited by Calliphona in the Canary Islands are compatible with a taxon cycle process. Our results have further implications for the higher level phylogeny of the subfamily Tettigoniinae and suggest that some of the tribes as currently delimited may not correspond to natural groups.     

Mitochondrial DNA phylogeny of Brimstone butterflies (genus Gonepteryx) from the Canary Islands and Madeira

Part of the mitochondrial Cytochrome Oxidase I gene was sequenced for seven species of Gonepteryx (Pieridac) butterflies. Four of the species are island endemics inhabiting the Canary Island archipelago and Madeira. The remaining three are European and African conspecifics. Sequence data were analysed phylogenetically by maximum parsimony and maximum likelihood methods. The resulting trees were used to deduce Canarian species' ancestry, sequential inter-island colonization and systematics. They suggest African ancestry for the Canary Island taxa and a colonization pattern, within the archipelago, compatible with the geological ages of the islands and other Canarian fauna: a colonization sequence from Africa to Tenerife and Gomera, followed by Tenerife to La Palma. The molecular phylogeny indicated that there are three Canarian endemics, G. cleobule, G. palmae and G. eversi from Tenerife, La Palma and Gomera, respectively.     

Population history of Berthelot's pipit: colonization, gene flow and morphological divergence in Macaronesia

The fauna of oceanic islands provide exceptional models with which to examine patterns of dispersal, isolation and diversification, from incipient speciation to species level radiations. Here, we investigate recent differentiation and microevolutionary change in Berthelot's pipit (Anthus berthelotii), an endemic bird species inhabiting three Atlantic archipelagos. Mitochondrial DNA sequence data and microsatellite markers were used to deduce probable colonization pathway, genetic differentiation, and gene flow among the 12 island populations. Phenotypic differentiation was investigated based on eight biologically important morphological traits. We found little mitochondrial DNA variability, with only one and four haplotypes for the control region and cytochrome b, respectively. However, microsatellite data indicated moderate population differentiation (FST=0.069) between the three archipelagos that were identified as genetically distinct units with limited gene flow. Both results, combined with the estimated time of divergence (2.5 millions years ago) from the Anthus campestris (the sister species), suggest that this species has only recently dispersed throughout these islands. The genetic relationships, patterns of allelic richness and exclusive alleles among populations suggest the species originally colonized the Canary Islands and only later spread from there to the Madeiran archipelago and Selvagen Islands. Differentiation has also occurred within archipelagos, although to a lesser degree. Gene flow was observed more among the eastern and central islands of the Canaries than between these and the western islands or the Madeiran Islands. Morphological differences were also more important between than within archipelagos. Concordance between morphological and genetic differentiation provided ambiguous results suggesting that genetic drift alone was not sufficient to explain phenotypic differentiation. The observed genetic and morphological differences may therefore be the result of differing patterns of selection pressures between populations, with Berthelot's pipit undergoing a process of incipient differentiation.     

Evolutionary dynamics of satellite DNA family PIM357 in species of the genus Pimelia (Tenebrionidae,Coleoptera)

A large number of repeats of a satellite DNA (stDNA) family have been cloned and sequenced from species and populations of the genus Pimelia (Tenebrionidae, Coleoptera). The beetles were collected in the Canary Islands, Morocco, the Iberian Peninsula, and the Balearic Islands in order to analyze the evolutionary forces and processes acting on abundant stDNAs conserved at the genus level. This repetitive family is composed of an abundant A-T-rich stDNA, with basic units of 357 bp. All the sequences obtained showed similarity to the 22 repeat units of the PIM357 stDNA family described previously for six Iberian Pimelia species (Pons et al. 1997 ). An analysis based on similarity shows the presence of three different groups of sequences clearly in accordance with their geographical origin. One is composed of satellite sequences from Iberian and Balearic species, a second group from the Moroccan taxa, whereas the third one is from the Pimelia species endemic to the Canary Islands. The latter group shows higher nucleotide diversities for their stDNA sequences and a lack of relationship between transition stages to fixation and sequence divergence. Phylogeographic data of Canarian Pimelia show that the PIM357 stDNA family has persisted for more than 8 Myr and could probably be traced to the origin of the lineage. The data suggest that distinct demographic and phylogenetic patterns related to the colonization of the volcanic Canarian island chain account for particular evolutionary dynamics of the repeat DNA family in this group.     

Multiple colonizations, in situ speciation, and volcanism-associated stepping-stone dispersals shaped the phylogeography of the Macaronesian red fescues (Festuca L., Gramineae)

Whereas examples of insular speciation within the endemic-rich Macaronesian hotspot flora have been documented, the phylogeography of recently evolved plants in the region has received little attention. The Macaronesian red fescues constitute a narrow and recent radiation of four closely related diploid species distributed in the Canary Islands (F. agustinii), Madeira (F. jubata), and the Azores (F. francoi and F. petraea), with a single extant relative distributed in mainland southwest Europe (F. rivularis). Bayesian structure and priority consensus tree approaches and population spatial correlations between genetic, geographical, and dispersal distances were used to elucidate the phylogeographical patterns of these grasses. Independent versus related origins and dispersal versus isolation by distance (IBD) hypotheses were tested to explain the genetic differentiation of species and populations, respectively. Genetic structure was found to be geographically distributed among the archipelagos and the islands endemics. The high number of shared AFLP fragments in all four species suggests a recent single origin from a continental Pliocene ancestor. However, the strong allelic structure detected among the Canarian, Madeiran, and Azorean endemics and the significant standardized residual values obtained from structured Bayesian analysis for pairwise related origin hypotheses strongly supported the existence of three independent continental-oceanic colonization events. The Canarian F. agustinii, the Madeiran F. jubata, and the two sister F. francoi and F. petraea Azorean species likely evolved from different continental founders in their respective archipelagos. Despite the short span of time elapsed since colonization, the two sympatric Azorean species probably diverged in situ, following ecological adaptation, from a common ancestor that arrived from the near mainland. Simple dispersal hypotheses explained most of the genetic variation at the species level better than IBD models. The optimal dispersal model for F. agustinii was a bidirectional centripetal stepping-stone colonization pattern, an eastern-to-western volcanism-associated dispersion was favored for F. francoi, whereas for the recently derived F. petraea a counterintuitive direction of colonization (west-to-east) was suggested. The population-based phylogeographical trends deduced from our study could be used as predictive models for other Macaronesian plant endemics with similar distribution areas and dispersal abilities.     

Diversification of the forest beetle genus Tarphius on the Canary Islands, and the evolutionary origins of island endemics

The flightless beetle genus Tarphius Erichson (Coleoptera: Colydiidae) is a distinctive element of the beetle fauna of the Canary Islands with 29 species distributed across the five western islands. The majority of Tarphius species are rare and intimately associated with the monteverde forest and only two species occur on more than one island. In this study we investigate the phylogeography of the Canary Island Tarphius, and their relationship to Tarphius from the more northerly archipelagos of Madeira and the Azores using maximum parsimony and Bayesian inference analysis of mitochondrial cytochrome oxidase I and II sequence data. We use geological datings for the Canary Islands, Azores, and Madeira to calibrate specific nodes of the tree for the estimation of divergence times using a penalized likelihood method. Data suggest that the Canary Island species assemblage is of some antiquity, however, much of this species diversity is relatively recent in origin. The phylogenetic relationships of species inhabiting the younger islands of El Hierro and La Palma indicate that colonization events between islands have probably been a significant factor in the evolutionary history of the Canary Island species assemblage. A comparison of molecular phylogenetic studies of arthropods on the Canary Islands suggests that, in the evolution of the arthropod species community of an island, the origin of endemic species is initially the result of colonizing lineages differentiating from their source populations. However, as an island matures a greater proportion of endemic species originate from intra-island speciation.     

Colonization and diversification: towards a phylogeographic synthesis for the Canary Islands

Recently, the Canary Islands have become a focus for studies of the colonization and the diversification of different organisms. Some authors have considered Canarian endemisms as relicts of Tertiary origin, but new molecular data suggest a general pattern of continental dispersion followed by in situ speciation. Recent phylogeographic studies are revealing variants of the simple stepping-stone colonization model that seems to hold for many Hawaiian groups. Many factors can generate deviations from such a pattern: the stochastic nature of colonization, competitive exclusion, phylogenetic constraints on adaptive evolution and extinction. An understanding of island colonization and diversification can best be developed from an ecosystem level synthesis as more data for the Canarian archipelago come to hand.     

Long-distance colonization and radiation in gekkonid lizards, Tarentola (Reptilia: Gekkonidae), revealed by mitochondrial DNA sequences

Morphological systematics makes it clear that many non-volant animal groups have undergone extensive transmarine dispersal with subsequent radiation in new, often island, areas. However, details of such events are often lacking. Here we use partial DNA sequences derived from the mitochondrial cytochrome b and 12S rRNA genes (up to 684 and 320 bp, respectively) to trace migration and speciation in Tarentola geckos, a primarily North African clade which has invaded many of the warmer islands in the North Atlantic Ocean. There were four main invasions of archipelagos presumably by rafting. (i) The subgenus Neotarentola reached Cuba up to 23 million years (Myr) ago, apparently via the North Equatorial current, a journey of at least 6000 km. (ii) The subgenus Tarentola invaded the eastern Canary Islands relatively recently covering a minimum of 120 km. (iii) The subgenus Makariogecko got to Gran Canaria and the western Canary Islands 7-17.5 Myr ago, either directly from the mainland or via the Selvages or the archipelago of Madeira, an excursion of 200-1200 km. (iv) A single species of Makariogecko from Gomera or Tenerife in the western Canaries made the 1400 km journey to the Cape Verde Islands tip to 7 Myr ago by way of the south-running Canary current. Many journeys have also occurred within archipelagos, a minimum of five taking place in the Canaries and perhaps 16 in the Cape Verde Islands. Occupation of the Cape Verde archipelago first involved an island in the northern group, perhaps São Nicolau, with subsequent spread to its close neighbours. The eastern and southern islands were colonized from these northern islands, at least two invasions widely separated in time being involved. While there are just three allopatric species of Makariogecko in the Canaries, the single invader of the Cape Verde Islands radiated into five, most of the islands being inhabited by two of these which differ in size. While size difference may possibly be a product of character displacement in the northern islands, taxa of different sizes reached the southern islands independently.     

Are plant communities on the Canary Islands resistant to plant invasion?

Oceanic islands are renowned for their unique flora and high levels of endemism. Native island plants, however, are imperilled by non-native species that can become invasive by outcompeting natives. The threat of native island assemblages generally increases with isolation and the number of endemics featured, but also with human-associated disturbance and land use. Based on this, the Canary Island native plant systems should be highly threatened by invasives, similar to other oceanic islands globally. However, Canarian native plant systems are only weakly infiltrated and are rarely directly threatened by invasive plants. Further, highly disturbed areas, usually among the first colonized by invasives on islands, are recolonized here by natives. Based on this, we postulate four hypotheses (climatic filter, well-preservation status, human legacy and permanent colonization) for explaining this unusual behaviour of plant systems on the Canary Islands, providing an opportunity to understand the drivers and processes behind invasion into plant communities on islands.     

A multi-gene approach reveals a complex evolutionary history in the Cyanistes species group

Quaternary climatic oscillations have been considered decisive in shaping much of the phylogeographic structure around the Mediterranean Basin. Within this paradigm, peripheral islands are usually considered as the endpoints of the colonization processes. Here, we use nuclear and mitochondrial markers to investigate the phylogeography of the blue tit complex (blue tit Cyanistes caeruleus, Canary blue tit C. teneriffae and azure tit C. cyanus), and assess the role of the Canary Islands for the geographic structuring of genetic variation. The Canary blue tit exhibits strong genetic differentiation within the Canary Islands and, in combination with other related continental species, provides an ideal model in which to examine recent differentiation within a closely related group of continental and oceanic island avian species. We analysed DNA sequences from 51 breeding populations and more than 400 individuals in the blue tit complex. Discrepancies in the nuclear and mitochondrial gene trees provided evidence of a complex evolutionary process around the Mediterranean Basin. Coalescent analyses revealed gene flow between C. caeruleus and C. teneriffae suggesting a dynamic process with multiple phases of colonization and geographic overlapping ranges. Microsatellite data indicated strong genetic differentiation among the Canary Islands and between the Canary archipelago and the close continental areas, indicating limited contemporary gene flow. Diversification of the blue tit complex is estimated to have started during the early Pliocene (≈ 5 Ma), coincident with the end of Messinian salinity crisis. Phylogenetic analyses indicated that the North African blue tit is derived from the Canary blue tits, a pattern is avian 'back colonization' that contrasts with more traditionally held views of islands being sinks rather than sources.     

Scrophularia arguta,a widespread annual plant in the Canary Islands: a single recent colonization event or a more complex phylogeographic pattern?

Many studies have addressed evolution and phylogeography of plant taxa in oceanic islands, but have primarily focused on endemics because of the assumption that in widespread taxa the absence of morphological differentiation between island and mainland populations is due to recent colonization. In this paper, we studied the phylogeography of Scrophularia arguta, a widespread annual species, in an attempt to determine the number and spatiotemporal origins of dispersal events to Canary Islands. Four different regions, ITS and ETS from nDNA and psbA‐trnH and psbJ‐petA from cpDNA, were used to date divergence events within S. arguta lineages and determine the phylogenetic relationships among populations. A haplotype network was obtained to elucidate the phylogenetic relationships among haplotypes. Our results support an ancient origin of S. arguta (Miocene) with expansion and genetic differentiation in the Pliocene coinciding with the aridification of northern Africa and the formation of the Mediterranean climate. Indeed, results indicate for Canary Islands three different events of colonization, including two ancient events that probably happened in the Pliocene and have originated the genetically most divergent populations into this species and, interestingly, a recent third event of colonization of Gran Canaria from mainland instead from the closest islands (Tenerife or Fuerteventura). In spite of the great genetic divergence among populations, it has not implied any morphological variation. Our work highlights the importance of nonendemic species to the genetic richness and conservation of island flora and the significance of the island populations of widespread taxa in the global biodiversity.     

Patterns of cladogenesis in the venomous marine gastropod genus Conus from the Cape Verde islands

Isolated oceanic archipelagos are excellent model systems to study speciation, biogeography, and evolutionary factors underlying the generation of biological diversity. Despite the wealth of studies documenting insular speciation, few of them focused on marine organisms. Here, we reconstruct phylogenetic relationships among species of the marine venomous gastropod genus Conus from the Cape Verde archipelago. This small island chain located in the Central Atlantic hosts 10% of the worldwide species diversity of Conus. Analyses were based on mtDNA sequences, and a novel nuclear marker, a megalin-like protein, member of the low-density lipoprotein receptor gene family. The inferred phylogeny recovered two well-defined clades within Conus. One includes Cape Verde endemic species with larger shells, known as the "venulatus" complex together with C. pulcher from the Canary Islands. The other is composed of Cape Verde endemic and West Africa and Canary Island "small" shelled species. In both clades, nonendemic Conus were resolved as sister groups of the Cape Verde endemics, respectively. Our results indicate that the ancestors of "small" and "large" shelled lineages independently colonized Cape Verde. The resulting biogeographical pattern shows the grouping of most Cape Verde endemics in monophyletic island assemblages. Statistical tests supported a recent radiation event within the "small shell" clade. Using a molecular clock, we estimated that the colonization of the islands by the "small" shelled species occurred relatively close to the origin of the islands whereas the arrival of "large" shelled Conus is more recent. Our results suggest that the main factor responsible for species diversity in the archipelago may be allopatric speciation promoted by the reduced dispersal capacity of nonplanktonic lecithotrophic larvae.     

The barriers to oceanic island radiation in bryophytes: insights from the phylogeography of the moss Grimmia montana

Aim In contrast to angiosperms, bryophytes do not appear to have radiated in Macaronesia and the western Mediterranean. We evaluate if: (1) the apparent lack of radiation in bryophytes reflects our failure to recognize cryptic endemic species; (2) bryophytes are characterized by extremely low evolutionary rates; or (3) bryophytes have a high dispersal ability, which prevents genetic isolation. Location Worldwide, with a special emphasis on Macaronesia and the western Mediterranean. Methods Three chloroplast regions were sequenced from samples of the moss Grimmia montana from its entire distribution range. Network analyses, Fst and Nst statistics were used to describe and interpret the phylogeographical signal in the data. Results Despite significant phylogeographical signal in the chloroplast genome, which demonstrates limits to gene flow at the continental scale, repeated sister group relationships observed among accessions from different geographical areas suggest recurrent colonization patterns. These observations are consistent with mounting evidence that intercontinental distributions exhibited by many bryophyte species result from long‐distance dispersal rather than continental drift. Madeiran and western Mediterranean island haplotypes are either shared by, or closely related to, European and North American ones. Fst values between Madeira, western Mediterranean islands, North America and Europe are not significantly different from zero, and suggest that Madeira and the south‐western Mediterranean are subject to strong transatlantic gene flow. By contrast, haplotypes found in the Canary Islands are shared or closely related to those of populations from south‐western Europe or southern Africa. Main conclusions Multiple origins and colonization events are not consistent with the hypothesis of a relictual origin of the Macaronesian moss flora. One possible reason for the failure of taxa that experienced multiple colonization events to radiate is niche pre‐emption. We suggest that strong gene flow, coupled with the occupancy of all suitable niches, either by earlier conspecific colonizers or by other species, could be the mechanism preventing island radiation in G. montana and other cryptogams with high long‐distance dispersal abilities.