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.     

Explosive radiation of Cape Verde Conus, a marine species flock

Nearly 50 species of the marine gastropod genus Conus are restricted to the Cape Verde archipelago. This unusual concentration of endemics within a single set of oceanic islands is extremely uncharacteristic of marine taxa. Here we used phylogenetic analyses of 90 Conus species, including 30 endemics from Cape Verde, to reveal the relationships and origins of the endemic Cape Verde Conus. Results show that these species group in two distinct clades and represent a marine species flock that is restricted to a very narrowly confined geographical area. Species' originations occurred in exceptionally limited parts of the archipelago and in some cases radiations took place solely within single islands. Finally, comparison of levels of divergence between Cape Verde endemics and other Conus species suggests that the radiation of Conus in Cape Verde occurred during the last few million years.     

Insight into an island radiation: the Tarentola geckos of the Cape Verde archipelago

Aim To reassess the relationships between Tarentola geckos from the Cape Verde Islands by including specimens from all islands in the range. To determine the variation within forms by sequencing over 400 specimens, thereby allowing the discovery of cryptic forms and resolving some of the issues raised previously. This extensive sampling was also used to shed light on distributions and to explain genetic diversity by comparing the ages and ecological and geological features of the islands (size, elevation and habitat diversity). Location The Cape Verde Islands: an oceanic archipelago belonging to the Macaronesian biogeographic region, located around 500 km off Senegal. Methods A total of 405 new specimens of Tarentola geckos were collected from nine islands with very different geological histories, topography, climate and habitats. Mitochondrial cytochrome b (cyt b) gene and 12S rRNA partial sequences were obtained and analysed using phylogenetic methods and networks to determine molecular diversity, demographic features and phylogeographic patterns. Results The phylogenetic relationships between all known forms of Cape Verdean Tarentola specimens were estimated for the first time, the relationships between new forms were assessed and previously hypothesized relationships were re‐examined. Despite the large sample size, low intraspecific diversity was found using a 303‐bp cyt b fragment. Star‐like haplotype networks and statistical tests suggest the past occurrence of a rapid demographic and geographical expansion over most of the islands. Genetic variability is positively correlated with size, elevation and habitat diversity of the islands, but is not linearly related to the age of the islands. Biogeographical patterns have, in general, high concordance with phylogenetic breaks and with the three eco‐geographical island groups. Volcanism and habitat diversity, both tightly linked with island ontogeny, as postulated by the general dynamic model of oceanic island biogeography, as well as present and historical size of the islands appear to be the main factors explaining the genetic diversity of this group. Main conclusions The Tarentola radiation was clarified and is clearly associated with the geological and ecological features of the islands. Two factors may account for the low intraspecific variation: (1) recent volcanic activity and high ecological stress, and (2) poor habitat diversity within some islands. More studies are needed to align taxonomy with phylogenetic relationships, whereas GIS modelling may help to predict precise species distributions.     

Systematics,biogeography and evolution of the endemic Hemidactylus geckos (Reptilia,Squamata, Gekkonidae) of the Cape Verde Islands: based on morphology and mitochondrial and nuclear DNA sequences

A total of 1854 bp of mitochondrial DNA (669 bp of cytochrome b (cyt b) and 386 bp of 12S rRNA), and 804 bp of a nuclear gene (RAG2) were investigated in endemic Hemidactylus from eight Cape Verde Islands, and used to explore their phylogeny, biogeography and evolution. Maximum‐likelihood, maximum‐parsimony and Bayesian analyses based on mtDNA revealed four well‐supported clades with uncorrected genetic divergences of 7.8–12.4% in the cyt b plus 12S rRNA genes, which were also supported by nuclear DNA. A population from the southern island of Fogo is the most divergent in both molecules and morphology and is described as Hemidactylus lopezjuradoi sp. n., and the populations on Sal and Boavista are also assigned species status as H. boavistensis. Although divergent in their DNA, the clade on S. Nicolau and that in the north‐western islands are morphologically similar and both are assigned to H. bouvieri for the present. Hemidactylus b. razoensis from Raso is genetically similar to H. b. bouvieri and differs only in its smaller body size. A molecular clock suggests that the ancestor of the endemic Hemidactylus of the Cape Verde Islands colonized the archipelago approximately 10 ± 2.48 Mya, perhaps reaching the north‐eastern islands first. The H. lopezjuradoi lineage separated soon after, and the north‐western islands were colonized progressively but slowly, S. Nicolau probably being reached first, then S. Vicente and islands on the same bank, and finally Sto. Antão, which is likely to have been colonized less than 1 Mya. Hemidactylus boavistensis is abundant on the arid islands where it occurs, but H. bouvieri appears to have been uncommon at least since it was described 130 years ago, and the same may be true of H. lopezjuradoi sp. n. The impact of introduced H. angulatus and H. mabouia on the endemic Hemidactylus of the Cape Verde Islands is not clear, but the discovery of substantial genetic diversity in endemic Cape Verde Hemidactylus means that the conservation requirements of the group should be reassessed.     

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.     

Shallow subtidal macroalgae in the North-eastern Atlantic archipelagos (Macaronesian region): a spatial approach to community structure

Shallow subtidal macroalgal communities in the North-eastern Atlantic archipelagos (Azores, Madeira, Canaries and Cape Verde) were studied in order to identify their spatial organization patterns and the main drivers of change. Fifteen islands and 145 sites across 15º of latitude and 2850 km were sampled. We found high spatial variability across the scales considered (archipelago, island and site). The structure of macroalgal communities differed among archipelagos, except between Madeira and the Canaries, which were similar. Across a latitudinal gradient, macroalgal communities in the Azores were clearly separated from the other archipelagos; communities in Madeira and the Canaries occupied an intermediate position, while those in Cape Verde appeared at the opposite end of the gradient. In the Azores, species with warm-temperate affinities dominated communities. Cape Verde communities were, in contrast, dominated by tropical taxa, whereas in the subtropical Canaries and Madeira there was a mixture of species with colder and warmer affinities. Apart from crustose coralline algae, the Dictyotales were the group with greatest cover; larger and longer-lived species were progressively replaced by short-lived species along a latitudinal gradient from north to south. The perennial species Zonaria tournefortii dominated the sea-bottom in the Azores, the semi-perennial Lophophora variegata in the Canaries, the filamentous algae in Madeira and the ephemeral Dictyota dichotoma in Cape Verde. We hypothesized that the differences among archipelagos could be explained by synergies between temperature and herbivory, which increased in diversity southwards, especially in Cape Verde. This was supported by the predominance of non-crustose macroalgae in the Azores and of crustose macroalgae in Cape Verde, as would be predicted from the greater herbivore activity. At the scale of islands and sites, the same set of environmental variables drove differences in macroalgal community structure across all the Macaronesian archipelagos.     

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.     

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.     

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.     

Weight–length relationships for 27 demersal fish species caught off the Cape Verde archipelago (eastern North Atlantic)

Weight–length relationships were estimated for 27 demersal fish species of the Cape Verde archipelago. Samples were collected in October and November 2000 using longline gear in depths up to 600 m, and occasionally to 1200 m. The b values were within expected ranges, varying between 2.406 and 3.761, except for Syacium micrurum (1.574) and Paraconger notialis (4.476), which presented strong allometries instigated through further analysis. This work revises previous literature in the archipelago and provides the first reference on weight–length parameters for 10 fish species worldwide and for 21 species of the Cape Verde archipelago.     

The morphology, distribution and conservation implications of introduced rats, Rattus spp. in the granitic Seychelles

There are no native land mammals in the Seychelles archipelago other than bats. Introduced rats have reduced the conservation value of most islands. This paper compares the results of rat‐trapping carried out on eight islands in the granitic Seychelles, between July 1999 and April 2000. Trapping was carried out in both the dry and wet seasons. Three introduced rodent species were caught, including two species of rat (ship rat Rattus rattus Linnaeus and Norway rat R. norvegicus Berkenhout), but only one Rattus species occurred on each island. Both rat species were smaller than European or Asian conspecifics, and there were variations in the size and appearance of rats on different islands. Inter‐island differences in size and pelage colour are discussed.     

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.     

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.     

IUCN Red List assessment of the Cape Verde endemic flora: towards a global strategy for plant conservation in Macaronesia

We review the conservation status and threats to the endemic vascular flora of the Cape Verde islands, mostly based on the past two decades of collecting, literature review and herbarium specimens. The application of IUCN Red List criteria and categories using RAMAS software reveals that 78% of the endemic plants are threatened (29.3% Critically Endangered, 41.3% Endangered, 7.6% Vulnerable). Most of these endemics have a limited geographical range, and half of them have Areas of Occupancy and Extents of Occurrence of < 20 and 200 km², respectively. Our data show that, over the last two decades, the Cape Verde vascular plants have become more threatened and their conservation status has declined, mostly as a consequence of the increase in exotic species, habitat degradation and human disturbance. This paper presents the first comprehensive IUCN Red List data review for the plants endemic to Cape Verde, thus providing an important step towards the recognition and conservation of its threatened endemic flora at the national and global level. It also fills a knowledge gap, as it represents the first thorough assessment of the conservation status of the entire endemic flora of a Macaronesian archipelago.     

Relationships and evolution of the North African geckos,Geckonia and Tarentola (Reptilia: Gekkonidae), based on mitochondrial and nuclear DNA sequences

Mitochondrial (cytochrome b and 12S rRNA) and nuclear (c-mos) genes, analyzed by a variety of methods, indicate that the distinctive northwest African gecko Geckonia chazaliae is a member of the Tarentola clade, being most closely related to the species of the western Canary and Cape Verde islands. Relationships in Tarentola as a whole are as follows: (T. americana ((T. mauritanica, T. angustimentalis) ((T. deserti, T. boehmei) ((T. b. boettgeri-South (T. b. boettgeri-North (T. b. bischoffi, T. b. hierrensis))) ((T. annularis, T. ephippiata) (Geckonia, T. delalandii, T. gomerensis, Cape Verde species)))))); nearly all nodes have high bootstrap support. Results confirm that T. americana of Cuba and the Bahamas separated at the most basal dichotomy of the phylogeny and give no positive support for the monophyly of the subgenera Tarentola s. str. and Makariogecko. The latter includes Geckonia and the subgenus Sahelogecko. Continental Tarentola appear to have invaded the Sahara desert from its northern edge. They have also colonized groups of Atlantic islands five times: a single invasion of the West Indies and three of the Canary islands, one of which then went on to invade the Cape Verde archipelago. The phylogeny corroborates anatomical evidence that the ground-dwelling Geckonia had a climbing ancestry, something that is paralleled in some southern African terrestrial gekkonids related to Pachydactylus. Distinctive derived features of Geckonia occur in other gekkonids that are ground dwelling in arid habitats and may be functionally related to this environment. The evolution of such features indicates that, although Tarentola is generally very uniform and may have been so for over 10 million years, this is not due to any overwhelming phylogenetic constraint. G. chazaliae should be included in Tarentola, as Tarentola chazaliae.     

Hybridization and population structure of the Culex pipiens complex in the islands of Macaronesia

The Culex pipiens complex includes two widespread mosquito vector species, Cx. pipiens and Cx. quinquefasciatus. The distribution of these species varies in latitude, with the former being present in temperate regions and the latter in tropical and subtropical regions. However, their distribution range overlaps in certain areas and interspecific hybridization has been documented. Genetic introgression between these species may have epidemiological repercussions for West Nile virus (WNV) transmission. Bayesian clustering analysis based on multilocus genotypes of 12 microsatellites was used to determine levels of hybridization between these two species in Macaronesian islands, the only contact zone described in West Africa. The distribution of the two species reflects both the islands' biogeography and historical aspects of human colonization. Madeira Island displayed a homogenous population of Cx. pipiens, whereas Cape Verde showed a more intriguing scenario with extensive hybridization. In the islands of Brava and Santiago, only Cx. quinquefasciatus was found, while in Fogo and Maio high hybrid rates (～40%) between the two species were detected. Within the admixed populations, second-generation hybrids (～50%) were identified suggesting a lack of isolation mechanisms. The observed levels of hybridization may locally potentiate the transmission to humans of zoonotic arboviruses such as WNV.     

Explanations for bird species range size: ecological correlates and phylogenetic effects in the Canary Islands

Aim To explore the determinants of island occupancy of 48 terrestrial bird species in an oceanic archipelago, accounting for ecological components while controlling for phylogenetic effects. Location The seven main islands of the Canary archipelago. Methods We obtained field data on population density, habitat breadth and landscape distribution in Tenerife, Fuerteventura and La Palma, aiming to sample all available habitats and the gradient of altitudes. In total, 1715 line transects of 0.5 km were carried out during the breeding season. We also reviewed the literature for data on occupancy, the distance between the Canary Islands and the nearest distribution border on the mainland, body size and endemicity of the 48 terrestrial bird species studied. Phylogenetic eigenvector regression was used to quantify (and to control for) the amount of phylogenetic signal. Results The two measurements of occupancy (number of occupied islands or 10 × 10 km UTM squares) were tightly correlated and produced very similar results. The occupancy of the terrestrial birds of the Canary Islands during the breeding season had a very low phylogenetic effect. Species with broader habitat breadth, stronger preferences for urban environments, smaller body size, and a lower degree of endemicity had a broader geographical distribution in the archipelago, occupying a larger number of islands and 10 × 10 UTM squares. Main conclusions The habitat‐generalist species with a tolerance for novel urban environments tend to be present on more islands and to occupy a greater area, whereas large‐sized species that are genetically differentiated within the islands are less widespread. Therefore, some properties of the ranges of these species are explicable from basic biological features. A positive relationship of range size with local abundance, previously shown in continental studies, was not found, probably because it relies on free dispersal on continuous landmasses, which may not be applicable on oceanic islands.     

Playback experiments indicate absence of vocal recognition among temporally and geographically separated populations of Madeiran Storm-petrels Oceanodroma castro

A number of lines of evidence suggest that temporally segregated sympatric populations of Madeiran Storm‐petrels Oceanodroma castro breeding in the Azores are reproductively isolated and morphologically and genetically distinct from each other. Within the Galapagos Islands, similar sympatric populations may also be isolated from each other, as individuals are not known to switch breeding seasons. The taxonomic relationships among populations of this species that are seasonally and spatially separated are unclear and in need of revision. In this study, playback experiments were used to determine the level of vocal response among prospecting Madeiran Storm‐petrels at colonies in the Azores, Galapagos and Cape Verde islands to recordings from different populations. Vocalizations of all populations studied here differ in their structural characteristics and in all but one case prospecting Storm‐petrels showed far greater response to playback of burrow calls from their own colony type than to calls recorded at other seasonally or geographically distinct colonies. Additionally, the level of response to foreign colony types was no different to playback of vocalizations of an unrelated control species present at the same location. Although not all combinations of geographical and seasonal populations could be examined, the finding that prospecting hot‐season (breeding April–August) Storm‐petrels in the Azores did not differ in their response level to playback from Azores cool‐season (breeding August–March) storm‐petrels and Cory's Shearwaters Calonectris diomedea is of particular significance and suggests the existence of a pre‐mating isolation mechanism that would prevent interbreeding between these two sympatric populations. Furthermore, Azores hot‐season Storm‐petrels showed a similar absence of response to playback from Galapagos dry‐season (May–July) populations, indicating that they are also taxonomically distant from this group. Madeiran Storm‐petrels in the Cape Verde islands showed a low response rate to Azores hot‐season vocalizations, which did not differ from the response to unrelated controls. These data provide further evidence that the hot‐season Azores population represents a distinct taxon that is reproductively isolated from the sympatrically breeding cool‐season population, as well as from more distant populations in the Cape Verde and Galapagos islands.     

Inventaire des moustiques (Diptera : Culicidae) des îles du sud-ouest de l’océan Indien,Madagascar excepté — Une revue critique

Inventory of the mosquitoes (Diptera: Culicidae) of the islands of southwestern Indian Ocean, Madagascar excluded – A Critical Review. The biodiversity of mosquitoes in the islands of southwestern Indian Ocean is the concern of numerous publications. Here, we propose a synthetic inventory and the analysis of the mosquito diversity, based on the available literature. A comprehensive annotated checklist of mosquito species has been recently published on Madagascar; this is the reason why this land is excluded from our work. Studied area encompasses 28 tropical islands in the southern hemisphere: 4 islands in the Comoros archipelago, 5 Scattered Islands (îles Éparses), 5 in Mascarene, 11 in the Seychelles and 3 in the Chagos archipelago. In total, the mosquito list presents 73 valid species, of which 10 are Anophelinae and 63 Culicinae. The number of species that are distributed in these islands only is 19, i.e. 26%, which is a remarkable level for endemism. The richness in mosquito species in these islands is analysed through several aspects including geography, local speciation and natural or human dissemination. This updated inventory increases by 33% the number of known species by regard to the previous inventory published by Julvez & Mouchet in 1994. The historical responsibility of humans in the introduction of new mosquito species in these islands is strongly documented. For instance, the species with the highest distribution among islands are Aedes aegypti, Ae. albopictus and Culex quinquefasciatus. The islands belong to the afrotropical biogeographic area and, logically, the majority (63%) of mosquito species present phylogenetic affinities with continental Africa and/or Madagascar; interestingly, the number of species present in these islands and in Madagascar but absent in continental Africa is higher than the number of species present in these islands and in continental Africa but absent in Madagascar (respectively 12 and 2 species). Thanks to valuable increase in the sampling effort, our knowledge of the culicidian fauna is increasing in these islands that constitute indisputably hotspots of biodiversity.     

Systematic study of the genusDelilia (Asteraceae,Heliantheae)

A taxonomic study of the genusDelilia is presented. Two species are recognized:D. biflora, a widespread weed of tropical and subtropical regions of the New World, extending from Mexico to northern Argentina and recently introduced in the Cape Verde Islands; andD. repens, endemic to the Galapagos islands. A key to the species, illustrations, distribution maps, and complete synonymy are presented.