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.     

A COMPARATIVE STUDY OF TEMPERATURE RESPONSES OF CARIBBEAN SEAWEEDS FROM DIFFERENT BIOGEOGRAPHIC GROUPS1

Temperature tolerances were determined for Caribbean isolates (total 31) of seaureds belonging to three distributional groups: 1) species confined to the tropical western Atlantic (Botryocladia spinulifera, Chamaedoris peniculum, Cladophoropsis sundanensis, Dictyopteris justii, Dictyurus occidentalis, Haloplegma duperreyi, and Heterosiphonia gibbesii); 2) amphi-Atlantic species with a (sub)tropical distribution that have their northern boundary in the eastern Atlantic at the tropical Cape Verde Islands (Bryothamnion triquetrum and Ceramium nitens) or the subtropical Canary Islands (Ceratodictyon intricatum, Coelothrix irregularis, Dictyopteris delicatula, Ernodesmis verticillata, and Lophocladia trichoclados; and 3) species with an am-phi-Atlantic tropical to warm-temperate distribution also occurring in the Mediterranean (Cladophoropsis membranacea, Digenea simplex, Microdictyon boergesenii, and Wurdemannia miniata). For some isolates, growth response curves and temperature requirements for reproduction were also determined. Growth occurred in the range (18)20–30° C with optimum growth rates at 25°–30°C, irrespective of distribution group. Reproduction generally occurred at (20)25°–30° C although there were some exceptions. Species were extremely stenothermal, with those restricted to the western Atlantic surviving a total range of only 10/13° C (between 18/20° and 30/33° C). Tolerance to high temperatures was correlated with vertical position in the iniertidal/subtidal zone rather than biogeography grouping. Species restricted to the subtidal were the least tolerant, with permanent survival at 30° C but not at 33°C. Tolerance to low temperatures was not different in subtidal and intertidal species but was significantly better in am phi-Atlantic than in western Atlantic species. In the former group, damage occurred at 15°–18° C but in the latter group at 18°-20° C. We propose that these differences in low-temperature tolerances in Caribbean populations of species from different distribution groups reflect adaptations to glacial cold-stress in the tropical eastern Atlantic and subsequent trans-Atlantic dispersal.     

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.     

Genetic relationships of the hydrocoral Millepora alcicornis and its symbionts within and between locations across the Atlantic

Although the hydrocoral Millepora alcicornis is a prominent and ecologically relevant amphi-Atlantic reef builder, little attention has been given to its endosymbionts which are also involved in the survival and adaptation success of the species in different environments. In this study, we resolve the genetic relationships between M. alcicornis and its symbionts (Symbiodiniaceae) within both sides and across the Atlantic. The COI and 16S-rDNA regions were selected for the host tissues, and the 23S-rDNA and ITS regions were chosen for the symbionts. Phylogenetic networks consistently showed that host populations from the eastern Atlantic archipelagos (Canary and Cape Verde Islands) were more related to western Atlantic populations than they were between them. However, results for Symbiodiniaceae species varied according to the molecular marker used. Samples from Mexico were grouped as Symbiodinium sp. (formerly Symbiodinium clade A) by both markers. Specimens from Puerto Rico were grouped as either Symbiodinium sp. or Breviolum sp. (formerly Symbiodinium clade B), according to the molecular marker used. Most samples from the eastern Atlantic were identified as Breviolum sp. by both markers, except for one sample from the Canary Islands and two samples from the Cape Verde Islands, which were identified as Cladocopium sp. (formerly Symbiodinium clade C) using ITS-rDNA. These results suggest that these two genera of Symbiodiniaceae may cohabit the same M. alcicornis colony. Because hydrocorals from the Canary Islands were phylogenetically related to the western Atlantic, but symbionts were more related to those of the Cape Verde Islands, the origin of the coral and its symbionts is probably different. This may be explained either by “horizontal” transmission, i.e. acquisition from the environment, or by a change in the dominant symbiont composition within the host. The flexibility of this hydrocoral to select symbionts, depending on environmental conditions, can provide new insight to understand how this coral may face ongoing climate change.

     

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.     

BIOGEOGRAPHY OF CLADOPHOROPSIS MEMBRANACEA (CHLOROPHYTA) BASED ON COMPARISONS OF NUCLEAR rDNA ITS SEQUENCES1

Nucleotides were compared at 988 sites, spanning both internal transcribed spacers (ITS1 and ITS2) of the nuclear ribosomal DNA, among 17 isolates of the green alga Cladophoropsis membranacea (Hofman Bang ex C. Agardh) Boergesen and two isolates of Struvea anastomosans (Harvey) Piccone and Grunow. Collections were made from Bonaire, Curaçao, St. Croix, the Canary Islands, the Cape Verde Islands, Mauritania, Syria, the Red Sea, Okinawa, and Hawaii. Two nucleotide substitutions were found between the sequenced coding regions of C. membranacea and S. anastomosans. Of the 720 nucleotides compared in ITS1 and ITS2, an average of 6.7% sequence divergence was found within C. membranacea and 17.4% between C. membranacea and S. anastomosans. Sequences were analyzed using maximum parsimony. Phylogenetic hypotheses were compared with the biogeographic hypothesis of an east-west Tethyan vicariance. Results of the study allow the identification of widely dispersed biogeographic populations, the identification of an underlying Tethyan imprint, and support for the hypothesis that dispersal has occurred between the Caribbean and the tropical eastern Atlantic. These results demonstrate the usefulness of ITS sequences at the sub-specific level in C. membranacea.     

Phylogeography of Ophioblennius: the role of ocean currents and geography in reef fish evolution

Many tropical reef fishes are divided into Atlantic and East Pacific taxa, placing similar species in two very different biogeographic regimes. The tropical Atlantic is a closed ocean basin with relatively stable currents, whereas the East Pacific is an open basin with unstable oceanic circulation. To assess how evolutionary processes are influenced by these differences in oceanography and geography, we analyze a 630-bp region of mitochondrial cytochrome b from 171 individuals in the blenniid genus Ophioblennius. Our results demonstrate deep genetic structuring in the Atlantic species, O. atlanticus, corresponding to recognized biogeographic provinces, with divergences of d = 5.2-12.7% among the Caribbean, Brazilian, St. Helena/Ascension Island, Gulf of Guinea, and Azores/Cape Verde regions. The Atlantic phylogeny is consistent with Pliocene dispersal from the western to eastern Atlantic, and the depth of these separations (along with prior morphological comparisons) may indicate previously unrecognized species. The eastern Pacific species, O. steindachneri, is characterized by markedly less structure than O. atlanticus, with shallow mitochondrial DNA lineages (dmax = 2.7%) and haplotype frequency shifts between locations in the Sea of Cortez, Pacific Panama, Clipperton Island, and the Galapagos Islands. No concordance between genetic structure and biogeographic provinces was found for O. steincdachneri. We attribute the phylogeographic pattern in O. atlanticus to dispersal during the reorganization of Atlantic circulation patterns that accompanied the shoaling of the Isthmus of Panama. The low degree of structure in the eastern Pacific is probably due to unstable circulation and linkage to the larger Pacific Ocean basin. The contrast in genetic signatures between Atlantic and eastern Pacific blennies demonstrates how differences in geology and oceanography have influenced evolutionary radiations within each region.     

On the occurrence of the hydrocoral Millepora (Hydrozoa: Milleporidae) in the subtropical eastern Atlantic (Canary Islands): is the colonization related to climatic events?

The occurrence of a hydrocoral of the genus Millepora has been recorded for the first time in the eastern subtropical Atlantic (Tenerife, Canary Islands), at a latitude of 11o N of its previously known northernmost limit of distribution in the Cape Verde Islands. The moderate development of the colonies, their fast growth rate and very restricted location indicate a recent colonization process, possibly related to an extreme climatic event that took place in the summer of 2004, adding to the rising seawater temperatures in the region during recent years.     

Genetic variation in epigenetic inheritance of ribosomal RNA gene methylation in Arabidopsis

We investigated the fidelity of epigenetic inheritance in crosses between three accessions of the flowering plant Arabidopsis thaliana (Canary Islands, Cape Verde Islands, and Columbia). Specifically, we examined the cytosine methylation content of the ribosomal RNA genes at the two nucleolus organizer regions (NOR2 and NOR4) in F1 and F2 hybrid individuals derived from reciprocal crosses between the high NOR methylation strain, Columbia, and the two other accessions, both of which have less NOR methylation. In crosses between the Columbia and Cape Verde Islands strains, the cytosine methylation content segregated as an additive Mendelian trait: the high NOR methylation state was tightly associated with the inheritance of the two Columbia-derived NOR loci. First-generation hybrid individuals between the Canary Islands and Columbia strains also showed a cytosine methylation content at the NORs intermediate between the parental values, consistent with the epigenetic inheritance of parental methylation patterns. Interestingly, mapping data from F2 individuals derived from a Canary Islands x Columbia cross revealed that NOR2 accounted for nearly all of the NOR methylation variation segregating in the population. NOR4 retains a significant effect on total NOR methylation content only through a complex epistatic interaction with NOR2. Our results indicate that the inheritance of differential cytosine methylation states at NOR loci can be modified by their genetic context, opening up the possibility of genetic dissection of epigenetic inheritance.     

Coconut varieties in Mexico

Cocos nucifera Linn, was introduced to Mexico by Spanish navigators in the middle of the 16th century, from Cape Verde and Santo Domingo to the Atlantic coast, and from the Solomon Islands and the Philippines to the Pacific coast. These populations remained isolated from one another for almost 500 years. Morphological studies on fruit characteristics show that coconuts from the two populations are contrastingforms. The Atlantic coast populations show a wild type fruit, with angular shape, thick husk, thick shell, thick endosperm and low water content. The Pacific populations have a domesticated type fruit, with fruit and nut more spherical, with a much thinner husk and shell, and higher water content. The higher variability observed in the Pacific coast raises the possibility of later introductions, also from the Philippines or other Pacific islands.     

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.     

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.     

Phylogeography of the Calonectris shearwaters using molecular and morphometric data

We investigated phylogenetic relationships and the biogeographic history of the Calonectris species complex, using both molecular and biometric data from one population of the Cape Verde shearwater Calonectris edwardsii (Cape Verde Islands), one from the streaked shearwater C. leucomelas (western Pacific Ocean) and 26 from Cory's shearwater populations distributed across the Atlantic (C. d. borealis) and the Mediterranean (C. d. diomedea). The streaked shearwater appeared as the most basal and distant clades, whereas the genetic divergences among the three main clades within the Palearctic were similar. Clock calibrations match the first speciation event within Calonectris to the Panama Isthmus formation, suggesting a vicariant scenario for the divergence of the Pacific and the Palearctic clades. The separation between the Atlantic and Mediterranean clades would have occurred in allopatry by range contraction followed by local adaptation during the major biogeographic events of the Pleistocene. The endemic form from Cape Verde probably evolved as a result of ecological divergence from the Mediterranean subspecies. Finally, one Mediterranean population (Almeria) was unexpectedly grouped into the Atlantic subspecies clade, both by genetic and by morphometric analyses, pointing out the Almeria-Oran oceanographic front (AOOF) as the actual divide between the two Cory's shearwater subspecies. Our results highlight the importance of oceanographic boundaries as potentially effective barriers shaping population and species phylogeographical structure in pelagic seabirds.     

Ovarian maturity,egg development,and offspring generation of the deep-water shrimp Plesionika edwardsii (Decapoda,Pandalidae) from three isolated populations in the eastern North Atlantic

Ovarian maturity, egg development and brood size were analysed for three isolated populations (Madeira, the Canary Islands and Cape Verde Islands) of Plesionika edwardsii (Decapoda, Pandalidae) in the eastern Atlantic. Multiple colour patterns were observed at the same ovarian maturity stage, which was verified histologically, invalidating the extensive use of ovarian colour as a maturity stage criterion. The physiological size at sexual maturity, based on the maturity of the ovaries, was higher in Madeira (carapace length of 19.73?mm) and decreased to the Cape Verdes (16.39?mm). Synchronic ovarian maturation was observed during the embryo incubation process, and ovigerous females bearing embryos at the final stage of development were found throughout the year. Females are multiple spawners during the reproductive season, after which the reproductive process ends and a rest period begins. The absence of females larger than the size at sexual maturity with ovaries in Stage 1, the incubation of embryos in the final developmental stages, suggests that the resting period begins with a process that reabsorbs the energy located in the ovaries and that the resting period occurs asynchronously in females in each of the studied populations. Embryo size was independent of female body size in the three populations studied, but increased with the developmental stage. A power equation was used to describe the relationship between brood size and female body size in the three areas studied. The mean number of external Stage I embryos carried by females decreased from Madeira (n?=?7868) to the Cape Verdes (n?=?3781), where less energy, in terms of the number of embryos and the size of the eggs, was invested in reproduction. Although female size decreases from north to south, the egg number was higher in Madeira than in the Cape Verdes for the same size range.     

Large scale population structure and gene flow in the planktonic developing periwinkle, Littorina striata, in Macaronesia (Mollusca: Gastropoda)

Allozymes were used to investigate the genetic structure of 42 populations of the planktonic developing, Macaronesian periwinkle Littorina striata, throughout its entire geographic range (Azores, Madeira, Canary Islands and Cape Verde Islands). This periwinkle is presumed to have a high dispersal and gene flow potential, because it has a planktonic development. It is therefore expected to show little population genetic differentiation. Indeed, based on Wright's hierarchical F-statistics, no significant genetic differentiation could be detected among populations, at any of the specified hierarchical levels (i.e. population, island, and archipelago). Nevertheless, the Cape Verde Islands seemed genetically more diverse (highest mean number of alleles per locus). The number of loci revealing a significant genetic heterogeneity increased with increasing distance between populations, while private alleles based gene flow (Nm) estimates also revealed a tendency towards a geographical pattern. The distribution of rare and private alleles, might account for these observations which suggested some geographical effect. Because of the low frequency at which these alleles occur, their influence on the genetic population structure is negligible, and not picked up by F-statistics.     

Hybridization studies in Bostrychia. 1: B. radicans (Rhodomelaceae, Rhodophyta) from Pacific and Atlantic North America

Bostrychia radicans(Montagne) Montagne is a pantropical/temperate red alga associated with mangroves and saltmarsh plants. Collections were made from a similar north-south geographic distribution along both the Pacific and Atlantic coasts of North America. Hybridization studies were performed with cultured isolates to assess the extent of interfertility and reproductive isolation along these two coastlines. All male and female gametophytes derived from single tetrasporophytes were intercompatible. Almost all isolates extending over 1500 km of coast line from northern Pacific Mexico are compatible, forming cystocarps that released viable carpospores. Even isolates which morphologically would be placed in two species [B. radicans and B. moritziana(Sender ex Kützing) J. Agardh], based on the presence or absence of monosiphonous branches, were capable of hybridizing. Crosses of isolates from the Atlantic USA showed a greater amount of incompatibility. Certain isolates were not compatible with any other isolates including isolates collected in close proximity (North Carolina isolates), while other isolates from the same locality were compatible (South Carolina). An isolate from South Carolina formed tetrasporophytes with isolates from Pacific Mexico but tetraspores were not viable. Certain incompatible crosses formed ‘pseudocystocarps’ but viable carposporophytes did not develop. Generalizations about reproductive isolation within a species must also consider differences between populations from different biogeographic regions that may reflect different paleoclimatological histories, founder effects and unique dispersal events.     

Genetic structure and history of populations of the deep-sea fish Helicolenus dactylopterus (Delaroche, 1809) inferred from mtDNA sequence analysis

Helicolenus dactylopterus is an Atlantic benthopelagic fish species inhabiting high-energy habitats on continental slopes, seamounts and islands. Partial sequences of the mitochondrial control region (D-loop) and cytochrome b (cyt b) were used to test the hypothesis that H. dactylopterus disperses between continental margin, island and seamount habitats on intraregional, regional and oceanic scales in the North Atlantic. Individuals were collected from five different geographical areas: Azores, Madeira, Portugal (Peniche), Cape Verde and the northwest Atlantic. D-loop (415 bp) and cyt b (423 bp) regions were partially sequenced for 208 and 212 individuals, respectively. Analysis of variation among mitochondrial DNA sequences based on pairwise F-statistics and AMOVA demonstrated marked genetic differentiation between populations in different geographical regions specifically the Mid-Atlantic Ridge (Azores)/northeast Atlantic (Portugal, Madeira) compared to populations around the Cape Verde Islands and in the northwest Atlantic. Some evidence of intraregional genetic differentiation between populations was found. Minimum-spanning network analysis revealed star-shaped patterns suggesting that populations had undergone expansion following bottlenecks and/or they have been colonized by jump dispersal events across large geographical distances along pathways of major ocean currents. Mismatch distribution analysis indicated that Azores and northwest Atlantic populations fitted a model of historical population expansion following a bottleneck/founder event estimated to be between 0.64 and 1.2 million years ago (Ma).     

A new species of the clingfish genus Apletodon (Teleostei: Gobiesocidae) from the Cape Verde Islands, Eastern Central Atlantic

The clingfish Apletodon barbatus sp. nov. is described on the basis of 22 specimens and color photos from Santiago and Sal Islands, Cape Verde Islands, eastern central Atlantic Ocean. The species is very small, apparently not exceeding 18 mm total length; it is characterized by having a conspicuous maxillary barbel in males, 4–5 incisors in the upper jaw, numerous brown spots on the head and body in males, and a double white spot near the anus. The new species is compared with other species of the genus; a key to the males of the 5 known species of the eastern Atlantic genus Apletodon is presented. A checklist is provided for the species of Apletodon and their synonyms. Several new records are included in the present paper: Apletodon dentatus and A. incognitus are recorded from the Canary Islands, and A. wirtzi is recorded from Cameroon.     

Population structure and conservation implications for the loggerhead sea turtle of the Cape Verde Islands

The Cape Verde Islands harbour the second largest nesting aggregation of the globally endangered loggerhead sea turtle in the Atlantic. To characterize the unknown genetic structure, connectivity, and demographic history of this population, we sequenced a segment of the mitochondrial (mt) DNA control region (380 bp, n = 186) and genotyped 12 microsatellite loci (n = 128) in females nesting at three islands of Cape Verde. No genetic differentiation in either haplotype or allele frequencies was found among the islands (mtDNA F _ST = 0.001, P > 0.02; nDNA F _ST = 0.001, P > 0.126). However, population pairwise comparisons of the mtDNA data revealed significant differences between Cape Verde and all previously sequenced Atlantic and Mediterranean rookeries (F _ST = 0.745; P < 0.000). Results of a mixed stock analysis of mtDNA data from 10 published oceanic feeding grounds showed that feeding grounds of the Madeira, Azores, and the Canary Islands, in the Atlantic Ocean, and Gimnesies, Pitiüses, and Andalusia, in the Mediterranean sea, are feeding grounds used by turtles born in Cape Verde, but that about 43% (±19%) of Cape Verde juveniles disperse to unknown areas. In a subset of samples (n = 145) we evaluated the utility of a longer segment (~760 bp) amplified by recently designed mtDNA control region primers for assessing the genetic structure of Atlantic loggerhead turtles. The analysis of the longer fragment revealed more variants overall than in the shorter segments. The genetic data presented here are likely to improve assignment and population genetic analyses, with significant conservation and research applications.     

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.