The genetic ghost of an invasion past: colonization and extinction revealed by historical hybridization in Senecio

Hybridization is an important evolutionary factor in the diversification of many plant and animal species. Of particular interest is that historical hybridization resulting in the origin of new species or introgressants has occurred between species now geographically separated by great distances. Here, we report that Senecio massaicus, a tetraploid species native to Morocco and the Canary Islands, contains genetic material of two distinct, geographically separated lineages: a Mediterranean lineage and a mainly southern African lineage. A time-calibrated internal transcribed spacer phylogeny indicates that the hybridization event took place up to 6.18 Ma. Because the southern African lineage has never been recorded from Morocco or the Canary Islands, we hypothesize that it reached this area in the distant past, but never became permanently established. Interestingly, the southern African lineage includes S. inaequidens, a highly invasive species that has recently become widespread throughout Europe and was introduced at the end of the 19th century as a 'wool alien'. Our results suggest that this more recent invasion of Europe by S. inaequidens represents the second arrival of this lineage into the region.     

Descriptions of Cosmocampus retropinnis sp. n., Minyichthys sentus sp. n. and Amphelikturus sp. (Pisces, Syngnathidae) from the Eastern Atlantic Region

Pelagic juvenile pipefishes, representing three genera previously unrecorded from the eastern Atlantic region, are reported from collections of the 1928–30 DANA expedition. Cosmocampus retropinnis sp. n., with dorsal-fin origin on tail and 6.75–8.0 subdorsal rings (S Morocco to Gambia), Minyichthys sentus sp. n., with patches of spinules on snout and 42 tail rings (W Mediterranean and Atlantic Ocean off N Morocco and Canary Is.), and Amphelikturus sp. (off Sierra Leone) are described and figured. The subgenus Minyichthys Herald and Randall is rediagnosed as a genus characterized, in part, by an 8-rayed caudal fin. A key is provided to the nine genera of Syngnathidae known from the eastern Atlantic region.     

Syntaxonomic analysis of theAndropogon-rich grasslands (Hyparrhenietalia hirtae) in the western Mediterranean region

Using the Braun-Blanquet approach, a syntaxonomical revision of the orderHyparrhenietalia hirtae in the western Mediterranean region was made. Various syntaxa were studied with relevés from the Iberian Peninsula, southern France, Sardinia, Sicily and adjacent islands, Balearic Islands, Canary Islands and Morocco. One alliance (Hyparrhenion hirtae), 8 associations (one of them new:Lotononido lupinifoliae-Hyparrhenietum sinaicae) and several subassociations and variants were distinguished. A complete classification of theHyparrhenietalia hirtae in the studied area with ecological and biogeographical diagnoses is given.     

Blitzkrieg in a Marine Invasion: Caulerpa racemosa var. cylindracea (Bryopsidales, Chlorophyta) Reaches the Canary Islands (North-East Atlantic)

On the basis of morphological and genetic studies (rDNA ITS1, 5.8S, ITS2, and a 18S rDNA intron), we confirm here that Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman et Boudouresque, a southwestern Australian taxon recently introduced into the Mediterranean Sea also occurs in the Canary Islands. This is the first report of C. racemosa var. cylindracea in the Atlantic. It was observed for the first time in the Canary Archipelago in 1997–1998. The speed and regional scale of expansion (north Atlantic Ocean and the Mediterranean Sea) of this invasive species appear to be among the most dramatic ever recorded. The possible outcome of this introduction in the Atlantic is discussed.     

Different fates of island brooms: contrasting evolution in Adenocarpus, Genista, and Teline (Genisteae, Fabaceae) in the Canary Islands and Madeira

Analysis of sequence data from the internal transcribed spacers (ITS) and 5.8S region of nuclear ribosomal DNA show that Canarian and Madeiran brooms (Genisteae) of the genera Teline, Adenocarpus, and Genista are related to Mediterranean species and not to species from adjacent parts of Morocco. Each separate colonization of the islands has resulted in contrasting patterns of adaptation and radiation. The genus Teline is polyphyletic, with both groups (the "T. monspessulana group" and the "T. linifolia group") separately nested within Genista. Genista benehoavensis (La Palma) and G. tenera (Madeira) form, with G. tinctoria of Europe, a single clade characterized by vestigially arillate seeds. The Canarian species of Adenocarpus have almost identical sequence to the Mediterranean A. complicatus and are likely to be the result of island speciation after a very recent colonization event. This Canarian/Mediterranean A. complicatus group is sister to the afrotropical montane A. mannii which is probably derived from an earlier colonization from the Mediterranean, possibly via the Red Sea hills. The independent colonization and subsequent radiation of the two Teline groups in the Canary Islands make an interesting comparison: the phylogenies both show geographical structuring, each with a central and western island division of taxa. Within the "T. monspessulana group" there is some evidence that both continental and Madeiran taxa could be derived from the Canary Islands, although it is likely that near contemporaneous speciation occurred via rapid colonization of the mainland and islands. The finding of two groups within Teline also has implications for patterns of hybridization in those parts of the world where Teline species are invasive; in California members of the T. monspessulana group hybridize readily, but no hybrids have been recorded with T. linifolia which has been introduced in the same areas.     

Biogeography and Genetic Structure in Populations of a Widespread Lichen (Parmelina tiliacea,Parmeliaceae, Ascomycota)

The genetic diversity and population structure of the foliose lichenized fungus Parmelina tiliacea has been analyzed through its geographical range, including samples from Macaronesia (Canary Islands), the Mediterranean, and Eurosiberia. DNA sequences from the nuclear ribosomal internal transcribed spacer, the mitochondrial large subunit ribosomal RNA gene, and the translation elongation factor 1-α were used as molecular markers. The haplotypes of the three markers and the molecular variance analyses of multilocus haplotypes showed the highest diversity in the Canary Islands, while restricted haplotypes occurred at high frequencies in Mediterranean coastal samples. The multilocus haplotypes formed three unevenly distributed clusters (clusters 1-3). In the Canary Islands all the haplotypes were present in a similar proportion, while the coastal Mediterranean sites had almost exclusively haplotypes of cluster 3; cluster 2 predominated in inland Mediterranean sites; and cluster 1 was more abundant in central and northern Europe (Eurosiberian area). The distribution of clusters is partially explained by climatic factors, and its interaction with local spatial structure, but much of the variation remains unexplained. The high frequency of individuals in the Canary Islands with haplotypes shared with other areas suggests that could be a refugium of genetic diversity, and the high frequency of individuals of the Mediterranean coastal sites with restricted haplotypes indicates that gene flow to contiguous areas may be restricted. This is significant for the selection of areas for conservation purposes, as those with most genetic variation may reflect historical factors and biological properties of the species.     

A tool set for description and mapping vegetation on protected natural areas: an example from the Canary Islands

The Canary Islands belong to Mediterranean basin hotspot, especially because their rich endemic flora. Nearly 40% of the total archipelago area is protected through protected natural area network. However, some plant communities show high degree of disturbance or they develop in unprotected zones. So, multivariate analysis combined with vegetation mapping can be useful for identify, describe and protect the main plant communities. These tools have been applied in Caldera de Taburiente national park (La Palma. Canary Islands). An actual vegetation map of this park is showed and its major natural vegetation units are described. These tools could be applied to other protected and unprotected areas, especially for planning, management and conservation purposes.     

Complex population genetic structure in the endemic Canary Island pine revealed using chloroplast microsatellite markers

The Canary archipelago, located on the northwestern Atlantic coast of Africa, is comprised of seven islands aligned from east to west, plus seven minor islets. All the islands were formed by volcanic eruptions and their geological history is well documented providing a historical framework to study colonization events. The Canary Island pine (Pinus canariensis C. Sm.), nowadays restricted to the westernmost Canary Islands (Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro), is considered an old (Lower Cretaceous) relic from an ancient Mediterranean evolutionary centre. Twenty seven chloroplast haplotypes were found in Canary Island pine but only one of them was common to all populations. The distribution of haplotypic variation in P. canariensis suggested the colonization of western Canary Islands from a single continental source located close to the Mediterranean Basin. Present-day populations of Canary Island pine retain levels of genetic diversity equivalent to those found in Mediterranean continental pine species, Pinus pinaster and Pinus halepensis. A hierarchical analysis of variance (AMOVA) showed high differentiation among populations within islands (approximately 19%) but no differentiation among islands. Simple differentiation models such as isolation by distance or stepping-stone colonization from older to younger islands were rejected based on product-moment correlations between pairwise genetic distances and both geographic distances and population-age divergences. However, the distribution of cpSSR diversity within the islands of Tenerife and Gran Canaria pointed towards the importance of the role played by regional Pliocene and Quaternary volcanic activity and long-distance gene flow in shaping the population genetic structure of the Canary Island pine. Therefore, conservation strategies at the population level are strongly recommended for this species.Communicated by D.B. NealeA. Gómez and S.C. González-Martínez as joint authors     

Mitochondrial differentiation and biogeography of Hyla meridionalis (Anura: Hylidae): an unusual phylogeographical pattern

Aim To study the patterns of genetic variation and the historical events and processes that influenced the distribution and intraspecific diversity in Hyla meridionalis Boettger, 1874. Location Hyla meridionalis is restricted to the western part of the Mediterranean region. In northern Africa it is present in Tunisia, Algeria and Morocco. In south‐western Europe it is found in the south of France, north‐western Italy and north‐eastern and south‐western Iberian Peninsula. There are also insular populations, as in the Canaries and Menorca. Methods Sampling included 112 individuals from 36 populations covering the range of the species. We used sequences of mitochondrial DNA Cytochrome Oxidase I (COI) for the phylogeographical analysis (841 bp) and COI plus a fragment including part of tRNA lysine, ATP synthase subunits 6 and 8 and part of Cytochrome Oxidase III for phylogenetic analyses (2441 bp). Phylogenetic analyses were performed with paup *4.0b10 (maximum likelihood, maximum parsimony) and Mr Bayes 3.0 (Bayesian analysis). Nested clade analysis was performed using tcs 1.18 and Geo Dis 2.2. A dispersal‐vicariant analysis was performed with diva 1.0 to generate hypotheses about the geographical distribution of ancestors. Results We found little genetic diversity within samples from Morocco, south‐western Europe and the Canary Islands, with three well‐differentiated clades. One is distributed in south‐western Iberia and the High Atlas, Anti‐Atlas and Massa River in Morocco. The second is restricted to the Medium Atlas Mountains. The third one is present in northern Morocco, north‐eastern Iberia, southern France and the Canaries. These three groups are also represented in the nested clade analysis. Sequences from Tunisian specimens are highly divergent from sequences of all other populations, suggesting that the split between the two lineages is ancient. diva analysis suggests that the ancestral distribution of the different lineages was restricted to Africa, and that an explanation of current distribution of the species requires three different dispersal events. Main conclusions Our results support the idea of a very recent colonization of south‐western Europe and the Canary Islands from Morocco. South‐western Europe has been colonized at least twice: once from northern Morocco probably to the Mediterranean coast of France and once from the western coast of Morocco to southern Iberia. Human transport is a likely explanation for at least one of these events. Within Morocco, the pattern of diversity is consistent with a model of mountain refugia during hyperarid periods within the Pleistocene. Evaluation of the phylogenetic relationships of Tunisian haplotypes will require an approach involving the other related hylid taxa in the area.     

Patterns of diversity of the north-eastern Atlantic blenniid fish fauna (Pisces: Blenniidae)

This paper presents an analysis of the distributional patterns of blenniids (Pisces: Blenniidae) in the north‐eastern Atlantic. Two peaks of species diversity were found, both in terms of number of species and number of endemics: one in the tropical African coast and another in the Mediterranean Sea. A cluster analysis of similarity values (Jaccard coefficient) among the eastern Atlantic zoogeographical areas, revealed the following groups: a north temperate group, a tropical group formed by the tropical African coast and Mauritania, another group formed by the islands of Cape Verde, a south temperate group (South Africa), and a southern Atlantic group formed by the islands of Ascension and St Helena. Within the north temperate group, the subgroups with higher similarities were: Azores and Madeira, Canary Islands and Morocco, and the Mediterranean and the Atlantic coast of the Iberian Peninsula. Based on affinity indices, the probable directions of faunal flows were inferred. The tropical coast of Africa and the Mediterranean emerged from this analysis as probable speciation centres of the north‐eastern Atlantic blenniid fauna. The Mediterranean may have also acted as a refuge during glacial periods.     

Nuclear ribosomal DNA and karyotypes indicate a NW African origin of South American Hypochaeris (Asteraceae, Cichorieae)

Hypochaeris has a disjunct distribution, with more than 15 species in the Mediterranean region, the Canary Islands, Europe, and Asia, and more than 40 species in South America. Previous studies have suggested that the New World taxa have evolved from ancestors similar to the central European H. maculata. Based on internal transcribed spacer (ITS) sequences and fluorescence in situ hybridization (FISH) with 5S and 18S-25S rDNA of the previously overlooked Hypochaeris angustifolia from Moyen Atlas, Morocco, we show that it is sister to the entire South American group. A biogeographic analysis supports the hypothesis of long-distance dispersal from NW Africa across the Atlantic Ocean for the origin of the South American taxa rather than migration from North America, through the Panamian land bridge, followed by subsequent extinction in North America. With the assumption of a molecular clock, the trans-Atlantic dispersal from NW Africa to South America is roughly estimated to have taken place during Pliocene or Pleistocene.     

A new endogean, anophthalmous species of Parazuphium Jeannel from Northern Morocco (Coleoptera, Carabidae), with new molecular data for the tribe Zuphiini

A new species of the genus Parazuphium (Coleoptera, Carabidae, Zuphiini), Parazuphium aguileraisp. n., is described from the Tingitan peninsula in North Morocco. The only known specimen was found under a large deeply buried boulder, and belongs to an anophthalmous, depigmented and flattened species. This is the second species of blind Parazuphium known so far, the other being Parazuphium feloi Machado 1998 from a lava tube in the Canary Islands. Molecular data of the only known Parazuphium aguilerai sp. n. specimen are provided, and a reconstructed phylogeny based on these molecular data confirms its inclusion inside Zuphiini within Harpalinae. Identification keys to the Mediterranean and Macaronesian species of Parazuphium are provided.     

Multilocus approach reveals an incipient differentiation process in the Stone-curlew, <Emphasis Type="Italic">Burhinus oedicnemus</Emphasis> around the Mediterranean basin

The Stone-curlew Burhinus oedicnemus, a steppe bird species, is mainly distributed in the Mediterranean and Macaronesian regions, which are considered hotspots of biodiversity with priority for animal and plant species richness conservation. In this study, we investigated the genetic diversity of the Stone-curlew in the Mediterranean basin and in the Canary Islands by applying a multilocus approach. We analysed mitochondrial and nuclear markers in order to evaluate the genetic structure and the congruence between morphological subspecies and geographic samples. We found a significant level of genetic differentiation between Mediterranean and Canary Island populations with all markers. Both in the Mediterranean basin and in the Canary Islands, we found a significant level of genetic diversity with nuclear markers only. We identified seven population groups, including insular populations. The four subspecies described for the Western Palaearctic were confirmed with some changes in distribution range. In spite of habitat fragmentation and negative population trend, the Stone-curlew showed a significant level of genetic diversity and gene flow among continental populations. However, islands constitute important reservoirs of genetic diversity and a potential for the evolution of the species.     

Parallel evolution of insular Olea europaea subspecies based on geographical structuring of plastid DNA variation and phenotypic similarity in leaf traits

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 (N_ST > G_ST = 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.     

Variation in wild populations of annual beet (Beta,Chenopodiaceae)

Among the morphological variation in wild annual populations of sect.Beta only tepal characters show a geographic pattern, and hence can be used to distinguish different taxa. Two morphological types correspond with taxa already described:B. macrocarpa (incl.B. bourgaei) has a Macaronesian and W. to E. Mediterranean andB. adanensis an E. Mediterranean distribution area. A third type in the Aegean region is not well known yet and possibly has to be included inB. macrocarpa. Both diploid and tetraploid (x = 9) cytotypes are found withinB. macrocarpa, the latter exclusively on the Canary Islands.     

Annotated check-list of the family Tachinidae (Diptera: Calyptratae: Oestroidea) in the Canary Islands

Summary

A commented check-list of the tachinid species inhabiting the Canary Islands (Spain) is provided, with special emphasis on hosts species and floral resources. Nine new local records are provided and Cylindromyia rufipes is reported for the first time for the Canary Islands, thus increasing the species number to 52. Zoogeographic analysis of the area revealed that there are differences in chorotype composition among islands, with the easternmost islands being richer in Mediterranean species and poorer in endemics. These differences may be due to ecological differences between the easternmost island and the remaining islands. Relevance regarding influence of tachinid as pollinators is discussed.     

Origin and evolution of the endemic Macaronesian Inuleae (Asteraceae): evidence from the internal transcribed spacers of nuclear ribosomal DNA

The tribe Inuleae (Asteraceae) has 10 species endemic to the Macaronesian islands, including the three endemic genera Allagopappus, Schizogyne, and Vierea. Phylogenetic analyses of DNA sequence data from the internal transcribed spacers (ITS) of the nuclear ribosomal DNA of 47 taxa were performed using all Macaronesian endemics and representative species from 21 of the 36 genera of the Inuleae. The resulting ITS phylogeny reveals that Allagopappus is sister to a large clade that contains all genera with a predominantly Mediterranean distribution. This finding suggests that Allagopappus may represent an ancient lineage that found refuge in the Canary Islands following the major climatic and/or geologic changes in the Mediterranean basin after the Tertiary. The Macaronesian endemic genus Schizogyne is sister to Limbarda from the Mediterranean. The third Macaronesian endemic genus, Vierea, is sister to Perralderia, which is restricted to Morocco and Algeria. Pulicaria canariensis is sister to P. mauritanica, a species endemic to Morocco and Algeria. In contrast, P. diffusa from the Cape Verde Islands is sister to a broadly distributed species, P. crispa, that occurs from North Africa to the Arabian peninsula. Based on the ITS data, the genera Blumea, Inula, and Pulicaria are not monophyletic. The ITS trees suggested that Blumea mollis belongs to the tribe Plucheeae, a finding that is congruent with recent morphological evidence. A possible southern African origin for the core of the Laurasian taxa of the Inuleae is also suggested.     

Phylogeography of the Atlanto-Mediterranean sea cucumber Holothuria (Holothuria) mammata: the combined effects of historical processes and current oceanographical pattern

We assessed the genetic structure of populations of the widely distributed sea cucumber Holothuria (Holothuria) mammata Grube, 1840, and investigated the effects of marine barriers to gene flow and historical processes. Several potential genetic breaks were considered, which would separate the Atlantic and Mediterranean basins, the isolated Macaronesian Islands from the other locations analysed, and the Western Mediterranean and Aegean Sea (Eastern Mediterranean). We analysed mitochondrial 16S and COI gene sequences from 177 individuals from four Atlantic locations and four Mediterranean locations. Haplotype diversity was high (H=0.9307 for 16S and 0.9203 for COI), and the haplotypes were closely related (π=0.0058 for 16S and 0.0071 for COI). The lowest genetic diversities were found in the Aegean Sea population. Our results showed that the COI gene was more variable and more useful for the detection of population structure than the 16S gene. The distribution of mtDNA haplotypes, the pairwise F(ST) values and the results of exact tests and amova revealed: (i) a significant genetic break between the population in the Aegean Sea and those in the other locations, as supported by both mitochondrial genes, and (ii) weak differentiation of the Canary and Azores Islands from the other populations; however, the populations from the Macaronesian Islands, Algarve and West Mediterranean could be considered to be a panmictic metapopulation. Isolation by distance was not identified in H. (H.) mammata. Historical events behind the observed findings, together with the current oceanographic patterns, were proposed and discussed as the main factors that determine the population structure and genetic signature of H. (H.) mammata.     

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.