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.     

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.     

Parallel gigantism and complex colonization patterns in the Cape Verde scincid lizards Mabuya and Macroscincus (Reptilia: Scincidae) revealed by mitochondrial DNA sequences

The scincid lizards of the Cape Verde islands comprise the extinct endemic giant Macroscincus coctei and at least five species of Mabuya, one of which, Mabuya vaillanti, also had populations with large body size. Phylogenetic analysis based on DNA sequences derived from the mitochondrial cytochrome b, cytochrome oxidase I and 12S rRNA genes (711, 498 and 378 base pairs (bp), respectively) corroborates morphological evidence that these species constitute a clade and that Macroscincus is unrelated to very large skinks in other areas. The relationships are ((M. vaillanti and Mabuya delalandii) (Mabuya spinalis and Macroscincus coctei (Mabuya fogoensis nicolauensis (Mabuya fogoensis antaoensis and Mabuya stangeri)))). The Cape Verde archipelago was colonized from West Africa, probably in the Late Miocene or Early Pliocene period. The north-eastern islands were probably occupied first, after which the ancestor of M. vaillanti and M. delalandii may have originated on Boavista, the ancestor of the latter species arriving on Santiago or Fogo later. The M. fogoensis--M. stangeri clade colonized the islands of Branco, Razo, Santa Luzia and São Vicente from São Nicolau and reached Santo Antão after this. Colonization of these northeastern islands was slow, perhaps because the recipient islands had not developed earlier or because colonization cut across the path of the Canary Current and the Northeast Trade Winds, the main dispersing agents in the region. Rapid extension of range into the southwestern islands occurred later in M. spinalis and then in M. vaillanti and M. delalandii. The long apparent delay between the origin of these species and their southwestern dispersal may have been because there were earlier colonizations of the southern islands which excluded later ones until the earlier inhabitants were exterminated by volcanic or climatic events. The evolution of large size in Macroscincus occurred in the northwestern islands and was paralleled in the eastern and southern islands by populations of M. vaillanti. Both cases of size increase in Cape Verde skinks were accompanied by the development of herbivory.     

A new species of Spauligodon (Nematoda: Oxyurida: Pharyngodonidae) in geckos from São Nicolau Island (Cape Verde) and its phylogenetic assessment

A new nematode species, Spauligodon nicolauensis n. sp., is described from geckos Tarentola bocagei and Tarentola nicolauensis on the island of S?o Nicolau, Cape Verde. The new nematode was found in the pellets obtained directly from the geckos in a non-invasive fashion, and its identity was assessed both at morphologic and genetic levels. The new species has morphological similarities with Spauligodon tarentolae Spaul, 1926, also parasitizing geckos from the Canary Islands. However, the male cloacal region in the new species is distinct, presenting a different shape of the caudal papillae. The overall resemblance probably resulted from colonization via descent from an ancestor of S. tarentolae carried by the ancestor of Cape Verde Tarentola. The analysis of nuclear DNA sequences confirms that the new species is phylogenetically distinct from all other Spauligodon species already analyzed, forming a group clearly separated from species parasitizing lacertid lizards. The COI genetic distance suggests that the S. nicolauensis n. sp. found in the 2 species of geckos in S?o Nicolau Island may have resulted from a host-switching event, when they came into contact after the unification of the island.     

Mitochondrial DNA patterns in the Macaronesia islands: Variation within and among archipelagos

Macaronesia covers four Atlantic archipelagos: the Azores, Madeira, the Canary Islands, and the Cape Verde islands. When discovered by Europeans in the 15th century, only the Canaries were inhabited. Historical reports highlight the impact of Iberians on settlement in Macaronesia. Although important differences in their settlement are documented, its influence on their genetic structures and relationships has yet to be ascertained. In this study, the hypervariable region I (HVRI) sequence and coding region polymorphisms of mitochondrial DNA (mtDNA) in 623 individuals from the Azores (120) and Canary Islands (503) were analyzed. Combined with published data, these give a total of 1,542 haplotypes from Macaronesia and 1,067 from the Iberian Peninsula. The results obtained indicate that Cape Verde is the most distinctive archipelago, with an mtDNA pool composed almost exclusively of African lineages. However, the other archipelagos present an mtDNA profile dominated by the presence of West‐Eurasian mtDNA haplogroups with African lineages present in varying proportions. Moreover, no signs of integration of typical Canarian U6 lineages in the other archipelagos were detected. The four Macaronesia archipelagos currently have differentiated genetic profiles, and the Azores present the highest intra‐archipelago differentiation and the lowest values of diversity. The analyses performed show that the present‐day genetic profile of the Macaronesian archipelagos was mainly determined by the initial process of settlement and further microdifferentiation probably as a consequence of the small population size of some islands. Moreover, contacts between archipelagos seem to have had a low impact on the mtDNA genetic pool of each archipelago. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

The Foraging Ecology of the Endangered Cape Verde Shearwater,a Sentinel Species for Marine Conservation off West Africa

Large Marine Ecosystems such as the Canary Current system off West Africa sustains high abundance of small pelagic prey, which attracts marine predators. Seabirds are top predators often used as biodiversity surrogates and sentinel species of the marine ecosystem health, thus frequently informing marine conservation planning. This study presents the first data on the spatial (GPS-loggers) and trophic (stable isotope analysis) ecology of a tropical seabird—the endangered Cape Verde shearwater Calonectris edwardsii–during both the incubation and the chick-rearing periods of two consecutive years. This information was related with marine environmental predictors (species distribution models), existent areas of conservation concern for seabirds (i.e. marine Important Bird Areas; marine IBAs) and threats to the marine environment in the West African areas heavily used by the shearwaters. There was an apparent inter-annual consistency on the spatial, foraging and trophic ecology of Cape Verde shearwater, but a strong alteration on the foraging strategies of adult breeders among breeding phases (i.e. from incubation to chick-rearing). During incubation, birds mostly targeted a discrete region off West Africa, known by its enhanced productivity profile and thus also highly exploited by international industrial fishery fleets. When chick-rearing, adults exploited the comparatively less productive tropical environment within the islands of Cape Verde, at relatively close distance from their breeding colony. The species enlarged its trophic niche and increased the trophic level of their prey from incubation to chick-rearing, likely to provision their chicks with a more diversified and better quality diet. There was a high overlap between the Cape Verde shearwaters foraging areas with those of European shearwater species that overwinter in this area and known areas of megafauna bycatch off West Africa, but very little overlap with existing Marine Important Bird Areas. Further investigation on the potential nefarious effects of fisheries on seabird communities exploiting the Canary Current system off West Africa is needed. Such negative effects could be alleviated or even dissipated if the ‘fisheries-conservation hotspots’ identified for the region, would be legislated as Marine Protected Areas.     

The Admixture Structure and Genetic Variation of the Archipelago of Cape Verde and Its Implications for Admixture Mapping Studies

Recently admixed populations offer unique opportunities for studying human history and for elucidating the genetic basis of complex traits that differ in prevalence between human populations. Historical records, classical protein markers, and preliminary genetic data indicate that the Cape Verde islands in West Africa are highly admixed and primarily descended from European males and African females. However, little is known about the variation in admixture levels, admixture dynamics and genetic diversity across the islands, or about the potential of Cape Verde for admixture mapping studies. We have performed a detailed analysis of phenotypic and genetic variation in Cape Verde based on objective skin color measurements, socio-economic status (SES) evaluations and data for 50 autosomal, 34 X-chromosome, and 21 non-recombinant Y-chromosome (NRY) markers in 845 individuals from six islands of the archipelago. We find extensive genetic admixture between European and African ancestral populations (mean West African ancestry = 0.57, sd = 0.08), with individual African ancestry proportions varying considerably among the islands. African ancestry proportions calculated with X and Y-chromosome markers confirm that the pattern of admixture has been sex-biased. The high-resolution NRY-STRs reveal additional patterns of variation among the islands that are most consistent with differentiation after admixture. The differences in the autosomal admixture proportions are clearly evident in the skin color distribution across the islands (Pearson r = 0.54, P-value<2e–16). Despite this strong correlation, there are significant interactions between SES and skin color that are independent of the relationship between skin color and genetic ancestry. The observed distributions of admixture, genetic variation and skin color and the relationship of skin color with SES relate to historical and social events taking place during the settlement history of Cape Verde, and have implications for the design of association studies using this population.     

An integrative taxonomic revision of the Tarentola geckos (Squamata,Phyllodactylidae) of the Cape Verde Islands

Recent phylogeographical analyses using mitochondrial DNA (mtDNA) sequences indicate that the Tarentola geckos from the Cape Verde archipelago originated from a propagule that dispersed from the Canary Islands approximately 7.7 Mya and that underwent a fast evolutionary radiation. Molecular analyses carried out to date clearly show some incongruences with the current taxonomy of Tarentola from the Cape Verde Islands, with some species being paraphyletic or polyphyletic, and several independently evolving lineages needing formal taxonomic recognition. The aim of this study was to clarify the systematics of this group to unravel its taxonomy by applying an integrative approach based on information from three independent sources: mtDNA, nuclear genes, and morphology. As a result of this taxonomic revision, two novel species for the islands of S. Nicolau and Fogo are described and eight subspecies are upgraded to species level. Moreover, an identification key for the genus Tarentola from the Cape Verde archipelago is presented. This study reconciles taxonomy and phylogeny in this group, provides a better understanding of diversity patterns, new insights on evolutionary hypotheses, and supports the basic framework for the future management and conservation of this unique reptile radiation. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 328–360.     

The apid cuckoo bees of the Cape Verde Islands (Hymenoptera, Apidae)

The apid cuckoo bees of the Cape Verde Islands (Republic of Cape Verde) are reviewed and five species recognized, representing two genera. The ammobatine genus Chiasmognathus Engel (Nomadinae: Ammobatini), a specialized lineage of cleptoparasites of nomioidine bees is recorded for the first time. Chiasmognathus batelkaisp. n. is distinguished from mainland African and Asian species. The genus Thyreus Panzer (Apinae: Melectini) is represented by four species - Thyreus denoliisp. n., Thyreus batelkaisp. n., Thyreus schwarzisp. n., and Thyreus aistleitnerisp. n. Previous records of Thyreus scutellaris (Fabricius) from the islands were based on misidentifications.     

Chloroplast DNA evidence for the roles of island colonization and extinction in Tolpis (Asteraceae: Lactuceae)

Tolpis consists of ～13 species native to Africa, Europe, and Macaronesia, with at least one species endemic to each of the four major archipelagos of the Azores, Madeira Islands, Canary Islands, and Cape Verde Islands. All but two of these species develop woody stems by maturity. Chloroplast DNA restriction site variation was analyzed for all species of Tolpis and four outgroups in order to understand the patterns of island colonization and evolution of woodiness in this genus. Parsimony analyses revealed a strongly supported monophyletic Tolpis. Within the genus, the following three well-supported groups were detected: all species from the Canary Islands and Cape Verde Islands, both Azorean species, and both continental species. The Canary Island/Cape Verde clade was sister to the two continental species, and the Azorean clade was sister to this group. The two Madeiran species of Tolpis occupied the basalmost positions within the genus. When biogeography was mapped onto this phylogeny, nine equally parsimonious reconstructions (five steps each) of dispersal history were detected, which fell into two groups: eight reconstructions implied that Tolpis colonized Madeira from the continent, followed by continental extinction and subsequent continental recolonization, while one reconstruction implied that Tolpis colonized Macaronesia four times. Two of the reconstructions involving continental extinction required the least amount of overall dispersal distance. The cpDNA phylogeny also suggests that woodiness arose in the common ancestor of all extant Tolpis, followed by two independent reversals to an herbaceous habit. Assuming that one of the eight reconstructions favoring continental extinction and recolonization is true, our results suggest that Tolpis may represent the first documented example of a woody plant group in Macaronesia that has recolonized the mainland in herbaceous form.     

Trophic habits of feral cats in the high mountain shrublands of the Macaronesian islands (NW Africa,Atlantic Ocean)

Feral cats Felis catus Linnaeus, 1758 have contributed to the extinction of numerous native species on islands, which are clearly sources of global biodiversity. We studied the diet of this introduced predator in the Madeira and Cape Verde archipelagos, which harbour important colonies of endangered seabirds in the high mountain habitats, and compared the results with those obtained in the same habitat in the Canary Islands, Macaronesian archipelago. On Madeira, 461 prey were identified from 143 scat groups. Mammals, overall mice, constituted the basic diet appearing in 95% of cat scats. On Fogo (Cape Verde), 657 prey items were obtained from 145 scats, and mammals were also the most important prey, reaching a frequency of occurrence of 88%. Although introduced mammals were the main prey category on all Macaronesian islands, we observed variation in feral cat diet among these islands. Birds were more frequently consumed on Madeira, lizards on Tenerife (Canaries) and invertebrates on Fogo. No specific differences were observed in relation to La Palma. We suggest that the diet composition on these islands varies according to the respective availability of the different prey types.     

EVOLUTIONARY AND ECOLOGICAL DIFFERENTIATION IN THE PANTROPICAL TO WARM-TEMPERATE SEAWEED DIGENEA SIMPLEX (RHODOPHYTA)1

Genetic differentiation among geographic isolates of the pantropical to warm-temperate red alga Digenea simplex (Wulfen) C. Agardh was investigated using random amplified polymorphic DNA (RAPD) markers, crossing studies, and temperature tolerances experiments. Eleven isolates representing populations from the Caribbean, eastern Atlantic, and Indo-West Pacific were compared. RAPD analysis clearly revealed an Indo-West Pacific group, a Caribbean/Cape Verde Islands group, and a Canary Islands group. Crossing studies showed different levels of inter fertility. In most crosses between Western Australian and Atlantic isolates, no hybrid tetrasporophytes were formed. In crosses between Caribbean and Cape Verde Islands isolates, tetrasporophytes developed, but the viability of tetraspores was reduced. Full sexual compatibility was observed among Cape Verde Islands isolates and among isolates from Bonaire. Temperature tolerance studies indicate that Pacific isolates have a broader temperature survival range than Atlantic isolates, which may be correlated to local temperature extremes. Despite the reduced level of sexual compatibility between Caribbean and Cape Verde Islands isolates, their shared position in the RAPD analysis and similar temperature responses suggest trans-Atlantic dispersal in the near geological past. In addition to their discrete position in the RAPD distance analysis, the Canary Islands isolates were significantly more cold-tolerant than the other Atlantic isolates. This finding is consistent with the hypothesis that the Canary Islands were recolonized from cold-adapted eastern Mediterranean populations after the last Pleistocene glaciation.     

PHYLOGEOGRAPHIC STUDIES IN THE TROPICAL SEAWEED CLADOPHOROPSIS MEMBRANACEA (CHLOROPHYTA,ULVOPHYCEAE) REVEAL A CRYPTIC SPECIES COMPLEX1

The seaweed Cladophoropsis membranacea (Hofman Bang ex. C. Agardh) Børgesen is a widely distributed species on coral reefs and along rocky coastlines throughout the tropics and subtropics. In a recent population‐level survey openface>1600 individuals with eight microsatellite loci, a number of isolates from biogeographically disjunct locations could not be amplified for any of the loci. Nonamplifiable and amplifiable isolates co‐occurred within the Canary Islands, Cape Verde Islands, and in the Caribbean. These unexpected results led to question whether or not C. membranacea is a single species. Phylogenetic relationships were evaluated using rDNA ITS1 and ITS2 sequence comparisons from 42 isolates sampled from a subset of 30 of the 66 locations. Four well‐supported clades were identified. Sequence divergence within clades was <1%, whereas between‐clade divergence was 2%–3%. Intraindividual variation was extremely low with no effects on the analysis. A strong, but imperfect, correspondence was found between ITS clades and amplifiable microsatellite loci. It is concluded that C. membranacea consists of three cryptic species. Using Pacific isolates as an outgroup, the most basal clade included the Central Canary Islands, Cape Verde, and Bonaire (Caribbean) isolates and thus spanned the widest latitude. Two derived sister clades consisted of a southern transtropical group stretching across the SE Caribbean to the Cape Verde Islands and African coast (but not the Canary Islands) and a NE‐Canary Island‐Mediterranean clade that also included the Red Sea. The detection of overlapping biogeographic distributions highlights the importance of ecotypic differentiation with respect to temperature and the importance of shifting sea surface isotherms that have driven periodic extinctions and recolonizations of the Canary Islands—a crossroads of marine floral exchange—since the last glacial maximum.     

Phylogeography of Cape Verde Island skinks (Mabuya)

The Cape Verde Islands are of volcanic origin with most having appeared between the early Miocene and mid-Pleistocene. They contain six known species of Mabuya skinks. Phylogeographical relationships within and among the relatively widespread taxa M. stangeri, M. spinalis and M. delalandii were inferred, based on approximately 1 kbp of the cytochrome b gene (mitochondrial DNA). Reciprocal monophyly of M. spinalis and M. stangeri was established, which may have arisen from an early Pliocene/late Miocene cladogenetic event. Considerable between-island sequence divergence was detected among M. spinalis, which appears to have colonized the older islands (Sal and Boavista) first. Much lower sequence divergence was found in M. delalandii, indicating a more recent range expansion. Here, evidence points to colonization of the younger islands of Brava and Fogo soon after appearance. There are similarities between some of the described patterns and those seen in lizards from the Canary Islands.     

Genetic diversity of the Macaronesian leafy liverwort Porella canariensis inferred from RAPD markers

Plant colonization of the North Atlantic raises the intriguing question of the relationships between extant island species with their continental counterparts (European, African, and American), which may provide clues to past geographic distribution and colonization history. It has been suggested that during past glaciations, many plant species with typical Mediterranean distributions survived in the Atlantic islands that belong to what is today known as Macronesia. We used random amplified polymorphic DNA (RAPD) markers to study 12 populations of the liverwort Porella canariensis partly covering its present-day distribution (Azores, Madeira, Canary and Cape Verde Islands, and Iberian Peninsula). Unweighted pair-group (UPGMA) and principal component (PCO) analyses showed a similar geographical pattern that suggested a close relationship between Iberian populations and those from the Canaries and Cape Verde Islands. Populations from Madeira had more genetic variation than those from the Azores, a result from either a richer diversity of habitats in Madeira, which prompted more population diversification, successive colonization waves from different origins, or an older colonization of Madeira. The data show that continuous patches of liverworts are often comprised of more than one individual. Finally, RAPDs can be used to investigate intraspecific diversity within a comparatively large geographic area and, with utmost care, can be used to infer a historic context to explain the patterns observed.     

Timing and tempo of early and successive adaptive radiations in Macaronesia

The flora of Macaronesia, which encompasses five Atlantic archipelagos (Azores, Canaries, Madeira, Cape Verde, and Salvage), is exceptionally rich and diverse. Spectacular radiation of numerous endemic plant groups has made the Macaronesian islands an outstanding area for studies of evolution and speciation. Despite intensive investigation in the last 15 years, absolute age and rate of diversification are poorly known for the flora of Macaronesia. Here we report molecular divergence estimates and rates of diversification for five representative, putative rapid radiations of monophyletic endemic plant lineages across the core eudicot clade of flowering plants. Three discrete windows of colonization during the Miocene and early Pliocene are suggested for these lineages, all of which are inferred to have had a single colonization event followed by rapid radiation. Subsequent inter-archipelago dispersal events into Madeira and the Cape Verdes took place very recently during the late Pliocene and Pleistocene after initial diversification on the Canary Islands. The tempo of adaptive radiations differs among the groups, but is relatively rapid compared to continental and other island radiations. Our results demonstrate that opportunity for island colonization and successful radiation may have been constrained to discrete time periods of profound climatic and geological changes in northern African and the Mediterranean.     

Timberline and alpine vegetation on the tropical and warm-temperate oceanic islands of the world: elevation, structure and floristics

In the oceans of the tropical and warm-temperate zone (40° N–40° S), only a small number of islands are high enough to show timberline and alpine vegetation. Excluding large islands with a more continental climate, only the following oceanic islands are relevant: Pico (Azores), Madeira, Tenerife, Gran Canaria and La Palma (Canary islands), Fogo (Cape Verde islands), Fernando Poo (Bioko) and Tristan da Cunha in the Atlantic Ocean, Réunion and Grande Comore (Ngazidja) in the Indian Ocean, Yakushima (Japan), Maui and Hawaii (Hawaiian islands), and Mas Afuera (Juan Fernandez islands) in the Pacific Ocean. Timberline and alpine vegetation exist here under a unique combination of a highly oceanic climate and a marked geographic isolation which contrasts with the tropical alpine vegetation in the extended mountains of South America, Africa and Southeast Asia.This review seeks to identify common physiognomic patterns in the high elevation vegetation that exist despite the fact that the islands belong to different floristic regions of the world. Based on the existing literature as well as personal observation, an overview of the elevation, physiognomy and floristics of the forest (and tree) line and the alpine vegetation on 15 island peaks is given.The forest line ecosystems are dominated either by conifers (Canary islands, Yakushima), heath woodland (Azores, Madeira, Réunion, Grande Comore, Fernando Poo) or broad-leaved trees (Hawaiian islands, Juan Fernandez islands, Tristan da Cunha). In the subalpine and alpine belts, dry sclerophyllous scrub occurs on island mountains that are exposed to the trade winds (Canary islands, Cape Verde islands, Hawaiian islands, Réunion, Grande Comore). These peaks are more or less arid above the forest line because a temperature inversion restricts the rise of humid air masses further upslope. In the summit regions of the remaining islands, which are located either in the wet equatorial and monsoonal regions or in the temperate westerly zones without an effective inversion layer, mesic to wet vegetation types (such as grassland, alpine heathland and fern scrub) are found.Compared to mountains at a similar latitude in continental areas, the forest line on the islands is found at 1000 to 2000 m lower elevations. The paper discusses four factors that are thought to contribute to this forest line depression: (1) drought on trade-wind exposed island peaks with stable temperature inversions, (2) the absense of well-adapted high-altitude tree species on isolated islands, (3) immaturity of volcanic soils, and (4) an only small mountain mass effect that influences the vertical temperature gradient.     

Complex estimates of evolutionary relationships in Tarentola mauritanica (Reptilia: Gekkonidae) derived from mitochondrial DNA sequences

Mitochondrial DNA (12S rRNA, 16S rRNA) sequences were analysed within Tarentola mauritanica and other selected species of Tarentola. Several highly genetically distinct lineages occur in North Africa, revealing phylogroups in southern and central Morocco, northern Morocco, Algeria, Tunisia, and Libya. A single haplotype characterizes populations across Spain, Portugal, Italy, Menorca, Crete, and Tunisia raising the possibility of an anthropogenic introduction followed by rapid population expansion throughout southern Europe. T. mauritanica is paraphyletic with respect to T. angustimentalis, a Canary islands endemic. The high genetic diversity observed across North Africa suggests T. mauritanica may represent a species complex.     

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.     

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.