Species richness and endemicity in the Spanish vascular flora

Data from an updated and revised checklist of the Spanish flora is analyzed. The Spanish vascular plant flora is composed of 204 families, 1433 genera and 7071 species. Floristic data are analysed by considering three regions: mainland Spain with 5984 species, the Balearic Islands with 1521 species and Canary Islands with 2066 species. Extinct species are included in the analysis, with an extinction rate of 0.35% of the flora. A total of 1488 endemic species are recognized, which account for 21% of the Spanish flora. The rate of endemism in the Canary Islands is 25.9%, considerably higher than for the Balearic Islands (6.9%) and for mainland Spain (13.8%). A list of the 35 strict endemic genera is compiled, of which 65.7% are Canarian, and another list of 27 subendemic genera of which 48.1% are also Canarian. An estimated 12% of the Spanish flora is non‐native, with large variation between the 20.7% of the Canary flora, 10.5% of mainland Spain and 9.7% of the Balearic Islands. Finally, the composition of the Spanish flora for large groups and families is ana lysed. The data show that species richness and endemicity rates have been overestimated by previous authors, and are similar to other Mediterranean countries. It is concluded that a complete revision of the Macaronesian flora is strongly needed to obtain an accurate comparison with the flora of other areas.     

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.     

Molecular systematics and biogeography of Resedaceae based on ITS and trnL-F sequences

The Resedaceae, containing 6 genera and ca. 85 species, are widely distributed in the Old World, with a major center of species diversity in the Mediterranean basin. Phylogenetic analyses of ITS and plastid trnL-trnF sequences of 66 species from all genera of the Resedaceae reveal (1) monophyly of the family, in congruence with preliminary phylogenetic studies; (2) molecular support for the traditional morphological subdivision of the Resedaceae into three tribes according to ovary and placentation types, and carpel number; (3) two monophyletic genera (Caylusea, Sesamoides), and one natural group (core Reseda), which includes the remaining four genera of the family (Ochradenus, Oligomeris, Randonia, Reseda); (4) a monophyletic origin for four of the six taxonomic sections recognized within Reseda (Leucoreseda, Luteola, Glaucoreseda, Phyteuma). Our results lead us to interpret an increment of the basic chromosome number in the family from x=5 to x=6 in at least two independent instances, and a broad representation of polyploids in multiple lineages across phylogenies, including association between octoploids and alien invasion in many parts of the world. Species diversity, endemism number, phylogenetic relationships and sequence divergence in Resedaceae suggest two major centers of differentiation, one in the western Mediterranean, and the other in the eastern Mediterranean and SW Asia. Two independent colonization events to the Canary Islands from Africa are indicated for the two Canarian Reseda endemics.     

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.     

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.     

Mitochondrial phylogeography of the long-eared bats (Plecotus) in the Mediterranean Palaearctic and Atlantic Islands

Long-eared bats of the genus Plecotus are widespread and common over most of the western Palaearctic. Based on recent molecular evidence, they proved to represent a complex of several cryptic species, with three new species being described from Europe in 2002. Evolutionary relationships among the different lineages are still fragmentary because of the limited geographic coverage of previous studies. Here we analyze Plecotus mitochondrial DNA sequences from the entire Mediterranean region and Atlantic Islands. Phylogenetic reconstructions group these western Palaearctic Plecotus into two major clades which split at least 5 Myr ago and that are each subdivided into further subgroups. An 'auritus group' includes the traditional P. auritus species and its sister taxon P. macrobullaris (=P. alpinus) plus related specimens from the Middle East. P. auritus and P. macrobullaris have broadly overlapping distributions in Europe, although the latter is apparently more restricted to mountain ranges. The other major clade, the 'austriacus group,' includes the European species P. austriacus and at least two other related taxa from North Africa (including P. teneriffae from the Canary Islands), the Balkans and Anatolia (P. kolombatovici). The sister species of this 'austriacus group' is P. balensis, an Ethiopian endemic. Phylogenetic reconstructions further suggest that P. austriacus reached Madeira during its relatively recent westward expansion through Europe, while the Canary Islands were colonized by a North African ancestor. Although colonization of the two groups of Atlantic Islands by Plecotus bats followed very distinct routes, neither involved lineages from the 'auritus group.' Furthermore, the Strait of Gibraltar perfectly segregates the distinct lineages, which confirms its key role as a geographic barrier. This study also stresses the biogeographical importance of the Mediterranean region, and particularly of North Africa, in understanding the evolution of the western Palaearctic biotas.     

Octopods of the Canary Islands. New records and biogeographic relationships

Two octopod species are reported from the Canary Islands (eastern Atlantic Ocean) for the first time: the deep sea four-horn octopus, Pteroctopus tetracirrhus (Delle Chiaje, 1830) and the gelatinous giant octopus, Haliphron atlanticus Steenstrup, 1861. Both female specimens were caught in Tenerife. Haliphron atlanticus is described from fresh remains found floating close to the southwest coast and the second species, P. tetracirrhus, is described from a specimen captured in a shrimp trap at 200 m depth on the southeastern coast of Tenerife. With these two additions the revised and updated list of octopod species of the Canary Islands now comprises eight families and 18 species, all of them incirrate octopods. The zoogeographic relationships of octopod species from other Atlantic regions, including the Mediterranean Sea, were studied. The likely directions of faunal flows were inferred based on affinity indices, showing that Mauritania could be the most probable source of the octopod species of the Canary Islands and the rest of the Macaronesian archipelagos.     

High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spermophora Hentz)

The six-eyed pholcid spiders of East Africa are revised. A new genus is recognized, Buitinga n. gen ., with 17 new species and three species transferred from Spermophora Hentz. The new genus is characterized by the presence of a scape on the epigynum. This scape may be straight or tightly curled up at rest, and is usually highly expandable. Seven additional African and Comoran species are newly described and tentatively assigned to Spermophora . A data matrix with 60 characters and 77 taxa (including 20 East African species and 25 additional Spermophora and ' Spermophora -like' species) is analysed cladistically. Buitinga is closer to the genera Paramicromerys Millot (endemic in Madagascar) and Spermophorides Wunderlich (Mediterranean and Canary Islands) as well as to several African and Comoran species tentatively assigned to Spermophora than to the type species of Spermophora . It is argued that current estimates of species numbers may be inaccurate and that pholcids may turn out to be one of the most diverse spider families.  © 2003 The Linnean Society of London. Zoological Journal of the Linnean Society, 2003, 137, 555−619     

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.     

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.     

Generic delimitation and phylogenetic relationships within the subtribe Chironiinae (Chironieae: Gentianaceae), with special reference to Centaurium: evidence from nrDNA and cpDNA sequences

To better understand the evolutionary history of the genus Centaurium and its relationship to other genera of the subtribe Chironiinae (Gentianaceae: Chironieae), molecular analyses were performed using 80 nuclear ribosomal ITS and 76 chloroplast trnLF (both the trnL UAA intron and the trnL-F spacer) sequences. In addition, morphological, palynological, and phytochemical characters were included to a combined data matrix to detect possible non-molecular synapomorphies. Phylogenetic reconstructions support the monophyly of the Chironiinae and an age estimate of ca. 22 million years for the subtribe. Conversely, both molecular data sets reveal a polyphyletic Centaurium, with four well-supported main clades hereafter treated as separate genera. The primarily Mediterranean Centaurium s.s. is closely related to southern African endemics Chironia and Orphium, and to the Chilean species Centaurium cachanlahuen. The resurrected Mexican and Central American genus Gyrandra is closely related to Sabatia (from eastern North America). Lastly, the monospecific genus Exaculum (Mediterranean) forms a monophyletic group together with the two new genera: Schenkia (Mediterranean and Australian species) and Zeltnera (all other indigenous American centauries). Several biogeographical patterns can be inferred for this group, supporting a Mediterranean origin followed by dispersals to (1) North America, Central America, and South America, (2) southern Africa (including the Cape region), and (3) Australia and Pacific Islands.     

Nuclear DNA amounts in Macaronesian angiosperms

Nuclear DNA contents for 104 Macaronesian angiosperms, with particular attention on Canary Islands endemics, were analysed using propidium iodide flow cytometry. Prime estimates for more than one-sixth of the whole Canarian endemic flora (including representatives of 11 endemic genera) were obtained. The resulting 1C DNA values ranged from 0.19 to 7.21 pg for Descurainia bourgeauana and Argyranthemum frutescens, respectively (about 38-fold difference). The majority of species, however, possessed (very) small genomes, with C-values <1.6 pg. The tendency towards small nuclear DNA contents and genome sizes was confirmed by comparing average values for Macaronesian and non-Macaronesian representatives of individual families, genera and major phylogenetic lineages. Our data support the hypothesis that the insular selection pressures in Macaronesia favour small C-values and genome sizes. Both positive and negative correlations between infrageneric nuclear DNA amount variation and environmental conditions on Tenerife were also found in several genera.     

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.     

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.     

Un'Ipotesi Sull'Evoluzione dei Limonium del Bacino del Mediterraneo

Abstract

A hypothesis on the evolution of Limonium species in the Mediterranean area. — The chromosomic somatic number of 27 species and 2 hybrids belonging to the genus Limonium (Plumbaginaceae) of several Mediterranean regions (Spain, France, Tuscany, Sardinia, Sicily) and Canarian Islands are here determined on wild populations. Keeping into account 10 data previously determined by other Authors, the chromosomic number is now known for more than 1/3 of the species occurring in West Europe and Mediterranean.

This allows to formulate a hypothesis on the group's evolution: from some diploid species living on inner rocks (Felsenheide) or sea cliffs and having a very restricted area, tetraploid races living on sandy shores had developped, which expanded afterwards on the whole Mediterranean area, and finally by hybridations giving rise to triploids.

Ecology and areal dimension of triploid species are generally intermediate between those of diploids and tetraploids; in some cases there is an apparent tendency to fragmentation into local forms, sometimes joined with apomictic reproduction.     

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.     

Molecular phylogenetics of the Macaronesian-endemic genus Bystropogon (Lamiaceae): palaeo-islands, ecological shifts and interisland colonizations

Abstract A molecular phylogenetic study of Bystropogon L'Her. (Lamiaceae) is presented. We performed a cladistic analysis of nucleotide sequences of the internal transcribed spacers (ITS), of the nuclear ribosomal DNA, and of the trnL gene and trnL-trnF intergenic spacer of the chloroplast DNA. Bystropogon odoratissimus is the only species endemic to the Canary Islands that occurs in the three palaeo-islands of Tenerife. This species is not part of an early diverging lineage of Bystropogon and we suggest that it has a recent origin. This phylogenetic pattern is followed by most of the species endemic to the palaeo-islands of Tenerife. The two sections currently recognized in Bystropogon form two monophyletic groups. Taxa belonging to the section Bystropogon clade show interisland colonization limited to the Canary Islands with ecological shifts among three ecological zones. Taxa from the section Canariense clade show interisland colonization both within the Canary Islands and between the Canary Islands and Madeira. Speciation events within this clade are mostly limited to the laurel forest. The genus has followed a colonization route from the Canaries towards Madeira. This route has also been followed by at least five other plant genera with species endemic to Macaronesia. Major incongruences were found between the current infrasectional classification and the molecular phylogeny, because the varieties of Bystropogon origanifolius and Bystropogon canariensis do not form two monophyletic groups. The widespread B. origanifolius appears as progenitor of the other species in section Bystropogon with a more restricted distribution.     

Distribution and biogeography of the Paradiaptominae (Copepoda: Calanoida: Diaptomidae)

The Paradiaptominae, a subfamily of the large Diaptomidae family, is a small group of freshwater calanoids of which 24 species in four genera have been described. They are endemic to Africa with the exception of four species, Metadiaptomus asiaticus (Asia and Mongolia), M. gauthieri (Africa and Madagascar), Paradiaptomus greeni (India and Sri Lanka) and Neolovenula alluaudi (Canary Islands, North Africa, Baltic and Mediterranean countries). The North African species Metadiaptomus chevreuxi extends its range into Iraq and Iran. The Paradiaptominae are widely distributed in semi-permanent water bodies in the drier areas of Africa although most individual species have a restricted distribution. Only a few species have the ability to colonise permanent waters and these species are usually located at higher altitudes. The Paradiaptominae have not been recorded from the wet equatorial lowland areas of Africa. It seems likely that they are Gondwanian in origin as evidenced by their distribution (Africa, Madagascar, India and Sri Lanka). Neolovenula alluaudi (Mediterranean) shares morphological characters tenuously linked to the other species in the group and may either have a different origin or have branched off at an early stage in evolution of the group.     

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     

Origin and evolution of the endemic Canary Island shrews (Mammalia: Soricidae)

Until recently the North Atlantic Islands were believed to house only mammals introduced by humans. Recent work has demonstrated that at least the Canary Islands house(d) a native mammal fauna. New data including chromosome numbers, genetic distances and analysis of vocalizations are given for the two extant shrew species, Crocidura canariensis and C. osorio , and their possible sister taxa are evaluated. Evidence is presented for the hypothesis that the two island species originated from two different lineages of the Palaearctic branch of the genus Crocidura. The data support the present status of the Canary Island shrews as local endemics of high conservation priority.