Phylogenetic relationships among the Ibero-African cobitids (Cobitis, cobitidae) based on cytochrome b sequence data

We estimated the phylogenetic relationships of all Ibero-African spined loaches of the genus Cobitis using the complete mitochondrial cytochrome b gene (1140bp). We analysed 93 individuals of seven cobitid species found in all the principal drainages of the Iberian Peninsula and North Africa. A molecular phylogeny was used to revise current systematics of the Ibero-African Cobitis species and to infer a biogeographical model for Cobitis in the Western Mediterranean area during the Cenozoic period. Phylogenetic analysis provided support for the monophyly of two mtDNA clades: Clade A or Italian Clade with the Italian species (C. bilineata, C. zanandreai), and Clade B or the Ibero-African Clade. The Ibero-African Clade included all species endemic for the Iberian Peninsula (C. vettonica, C. calderoni, and C. paludica) and North Africa (C. maroccana). The species C. paludica does not constitute a natural group, and could be divided into at least four monophyletic mtDNA lineages with moderate to high bootstrap values and posterior probability support. Phylogenetic relationships of the Ibero-African species were not resolved satisfactorily, but in all analyses C. calderoni from Northern Iberian Peninsula was basal. We have re-calibrated a molecular clock for the genus Cobitis (0.68% per million year by pairwise) using populations inhabiting both sides of the Gibraltar Strait. Application of this Cobitis mtDNA clock provides evidence that the Messinian salinity crisis played a primary role in the diversification of some Ibero-African cobitid species. The basal polytomies of the Ibero-African Clade might suggest that all mtDNA lineages diversified rapidly.     

Taxonomic study of the Iberian Cobitis (Osteichthyes, Cobitidae), with description of a new species

Morphometric and electrophoretic variation are analysed among Iberian and African populations of Cobitis (Cobitidae). Three discrete morphotypes are distinguished, showing a remarkable genetic differentiation among them. Each of the three morphotypes is judged to represent a species, one being described as a new species. Both approaches agree on the separate status of the Iberian populations from the Alagón drainage, Tagus basin of west central Spain. This population is described as a new species, Cobitis vettonica sp. nov., a sexually dimorphic spined loach was assigned to the subgenus Iberocobitis Bacescu, 1961. Morphometric and electrophoretic data placed this new species phenetically close to Cobitis maroccana Pellegrin, 1929 and Cobitis pahdica De Buen, 1930, but differs from these species in overall shape, male coloration pattern, shorter barbels and fins, posterior lamina of the cleithrum, seven branched rays on the dorsal fin and in having a unique alleles at sAAT-2*, EST-1* and MPI* loci. The new species range described in this paper is more restricted than that of C. pahdica. Both inhabit the Tagus basin but have not been reported to be sympatric.     

Phylogenetic relationships and historical biogeography of spined loaches (Cobitidae, Cobitis and Sabanejewia) as indicated by variability of mitochondrial DNA

For sequence analysis of the 12S rRNA gene in spined loaches, specimens of the following taxa were used: Sabanejewia balcanica, Cobitis paludica, C. bilineata, C. fahireae, C. elazigensis, C. elongata , two different sub-populations of C. taenia and four different sub-populations of C. turcica . Phylogenetic relationships among taxa were estimated using parsimony, neighbor-joining, and maximum likelihood algorithms. The calculation of the transition-tranversion ratio indicated that the taxa analysed were rather distantly related. Our analyses using the genus Sabanejewia as an outgroup suggest that C. bilineata is separated from the subgenus Cobitis sensu stricto and placed together with C. elongata as basal to all other species of the genus Cobitis . Support for the placement of C. paludica as basal to both the subgenera Bicanestrinia and Cobitis s. s. (without C. bilineata ) is given by outcomes from three independent methods of phylogenetic reconstruction. A sister-taxon relationship for the subgenera Bicanestrinia and Cobitis s. s. without C. bilineata was consistently found. C. elazigensis was closely related to the population of C. turcica from Lake Beysehir in Turkey. Both formed a sister-group to the remaining populations of C. turcica , while C. fahireae was basal to C. taenia . A molecular clock was calculated based on sequence divergence values and palaeogeographical data. This suggests that different historical colonization routes must have been used by different clades of spined loaches.     

Age, growth and reproduction of Cobitis paludica in a seasonal stream

In the Iberian endemic loach Cobitis paludica , in a small seasonal tributary of the Guadalquivir River, 65–70% of the total growth in length occurred in the first year of life. The maximum ages observed were 3+ years in males and 4+ years in females. Both sexes matured at the beginning of their second year of life. There was a significant difference from 1: 1 in the overall sex ratio of 412 males to 674 females. Spawning began in late March and ended in July. Cobitis paludica is a multiple spawner that releases a minimum of two batches of eggs per female each year. Once spawning had started, however, there was no recruitment from the stock of transparent oocytes to the mature stock. The number of eggs decreased from the first batch to the second, but there was no significant difference in the mean egg diameter between batches. Compared with other populations of this species, the present population, from a low latitude, is characterized by a low number of age groups, fast growth, early maturity and high fecundity in multiple spawnings. These life-history characteristics are typical of species in unstable environments, where adult mortality is high, variable or unpredictable.     

Cytotaxonomic studies in the sections Spirostachyae (Drejer) Bailey and Ceratocystis Dumort. of the genus Carex L. (Gyperaceae), with special reference to Iberian and North African taxa

LUCEÑO, M. & CASTROVIEJO, S., 1993. Cytotaxonomic studies in the sections Spirostachyae (Dreyer) Bailey and Ceratocystis Dumort of the genus Carex L. (Cyperaceae), with special reference to the Iberian and North African taxa. Karyological studies were carried out on Iberian and North African populations of all species included in sections Spirostachyae and Ceratocystis of the genus Carex , with observations made on meiotic divisions in one to seven individuals per population. In the light of the karyological data we present hypotheses of the most probable phylogeny of the section Spirostachyae , and of the relationships of the sections Spirostachyae, Ceratocystis and Elatae. Carex laevigata and C. camposii are transferred from section Elatae to section Spirostachyae.     

Colonization patterns and genetic structure of peripheral populations of the trumpeter finch (Bucanetes githagineus) from Northwest Africa, the Canary Islands and the Iberian Peninsula

Aim This paper has three aims: (1) to reconstruct the colonization history of two peripheral populations of the trumpeter finch (Bucanetes githagineus) presumably originating from the same source, one the result of an ancient expansion process and the other recently established and still expanding; (2) to estimate the importance of key events, such as past and current gene flow and bottlenecks, in both expansion processes and their contribution to the present population structure and genetic diversity; and (3) to find out whether two peripheral populations that established at widely differing times also differ in terms of genetic diversity. Location Northwest Africa (assumed source population), Canary Islands (long‐established peripheral) and south‐eastern Iberian Peninsula (recently established peripheral). Methods Bayesian analysis of population structure, individual assignment tests, F‐statistics, maximum likelihood migration estimates, genetic diversity indices and bottleneck tests were calculated with microsatellite data from 194 trumpeter finches from five breeding and two seasonal non‐breeding sites. Results Our data support the existence of two subpopulations (Canary Island and Ibero‐African) as the most likely population structure. Seasonal sites in the Iberian Peninsula had the highest percentage of birds assigned to other, mainly Iberian, sites. Pairwise F_ST values showed that the Canary Island localities were very similar to each other, but differed from the rest. Gene flow estimates within subpopulations were only slightly higher in the Canary Island population than in the Ibero‐African one. Gene diversity indices were similar at all localities. Canary Island sites show evidence of bottlenecks, whereas the Ibero‐African sites do not. Main conclusions Our data show that, at present, birds from the Canary Islands are genetically differentiated from those in North Africa and continental Spain. We could not unequivocally confirm the African origin of Canary populations because the contrary is also plausible. The Iberian Peninsula seems to have repeatedly received individuals from North Africa, which would have led to the relatively high genetic diversity found in these recently established localities and prevented bottlenecks. Movements of individuals towards sites outside their current range during the non‐breeding season are likely to precede the establishment of new breeding sites at the periphery of the distribution range.     

Spatio-temporal genetic variation in sympatric and allopatric Mediterranean Cicada species (Hemiptera, Cicadidae)

Cicada barbara (Stål) and Cicada orni L. are two Mediterranean cicadas, very similar in morphology, that produce distinct acoustic mating signals and that have partially overlapping distribution ranges in the Iberian Peninsula, occurring in sympatry in several locations. In the present study, six microsatellite loci were analysed in C. barbara , four of which were also analysed in C. orni . Geographical and temporal genetic variation in these species was studied. No evidence of hybridization was found, enabling us to infer that the isolating barriers between these species are efficient. Partitioning of geographic variation in each species, revealed the following patterns: Iberian Peninsula and Northwestern African populations of C. barbara showed higher differentiation between than within each region, supporting C. barbara subspecific taxonomy ( C. barbara lusitanica in the Iberian Peninsula and C. barbara barbara in Northwestern Africa) and highlighting isolation coincident with the presence of physical barriers to gene-flow; differentiation between populations of C. orni from both sides of the Pyrenees was very low, and this mountain range may not constitute a significant barrier for the dispersal of this species; Greek populations of C. orni were found to be highly differentiated from Western European populations; and no pattern of isolation-by-distance was found in either species within the Iberian Peninsula.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 249–265.     

Genetic and phenotypic variation among geographically isolated populations of the globally threatened Dupont's lark Chersophilus duponti

Identifying genetically and phenotypically distinct populations of threatened species is critical if we are to delineate appropriate plans for their conservation. We conducted an integrated analysis of population genetic structure, historical demographic events, current gene flow (all based on mtDNA sequences) and morphological variation of three geographically separated groups of populations of Dupont's lark Chersophilus duponti, located in the Iberian Peninsula (three populations), Morocco (two populations), and Tunisia (one population). Unusually, this lark species is the only one among the genus Chersophilus. Our results revealed the early historical divergence of an eastern Dupont's lark lineage (in Tunisia) and a western lineage (in Morocco and Spain), consistent with subspecies taxonomy and distribution. The western lineage subsequently split into two lineages, following the isolation of Iberian and African populations. Such pattern of historical differentiation caused great population genetic structure, with differences among geographic areas explaining more than 80% of total genetic variation. Mismatch distributions and coalescent estimates of divergence time showed that lineage divergence was associated with sudden population expansion events, which apparently took place during the last glaciation, when steppe habitats were widespread across the Mediterranean region. Extant populations from different geographic areas hardly shared any haplotype (only one out of 16 ND2 haplotypes was shared by Tunisian and Moroccan Dupont's larks), and consequently gene flow between geographic areas was found to be virtually absent. Apart from showing great genetic differentiation, Dupont's larks from different geographic areas were morphologically distinct, showing substantial variation in body size and feeding-related traits (length of feet and bill). We conclude that Dupont's lark populations isolated in the Iberian Peninsula, Morocco, and Tunisia are distinct evolutionary entities and should be considered as such in conservation plans. Such circumstance sets a daunting conservation challenge that exemplifies the need of incorporating knowledge of historical processes to our general understanding of the demography of threatened species.     

Population structure of Cicada barbara Stål (Hemiptera, Cicadoidea) from the Iberian Peninsula and Morocco based on mitochondrial DNA analysis

We assess the genetic history and population structure of Cicada barbara in Morocco and the Iberian Peninsula, based on analysis of the mitochondrial cytochrome b gene. The divergence between Morocco and the Iberian Peninsula populations was strongly corroborated by the molecular data, suggesting genetically isolated populations with a low level of gene flow. The Ceuta population from Spanish North Africa was more similar to the Iberian populations than the surrounding Moroccan populations, suggesting that the Strait of Gibraltar has not been acting as a strict barrier to dispersal while the Rif Mountains have. The Iberian Peninsula specimens showed a signature of demographic expansion before that which occurred in Morocco, but some of the assumptions related to the demographic parameters should be considered with caution due to the small genetic variation found. The high haplotype diversity found in Morocco implies higher demographic stability than in the Iberian Peninsula populations. These results do not, however, suggest a Moroccan origin for Iberian cicadas; but the most northwest region in Africa, such as Ceuta, might have acted as a southern refuge for Iberian cicadas during the most severe climatic conditions, from where they could expand north when climate improved. The separation of two subspecies within C. barbara (C. barbara lusitanica and C. barbara barbara) finds support with these results.     

Evidence of connectivity between continental and differentiated insular populations in a highly mobile species

Aim Genetically differentiated insular populations are candidates for independent units for conservation. However, occasional immigration to reduced island populations may occur and potentially have important consequences in their future viability and evolutionary potential. In this study, we investigate the conservation implications of population structure and connectivity of insular and continental populations of a migratory raptor as determined using genetic tools and satellite tracking. Location Western European populations in the Iberian Peninsula and two insular populations in the Mediterranean Sea (Balearic Islands) and Atlantic Ocean (Canary Islands). Methods We genotyped 22 microsatellite loci in 96 Egyptian vultures (Neophron percnopterus) from the Iberian Peninsula, 36 from Menorca (Balearic archipelago) and 242 (85% of the current population) from Fuerteventura (Canary Islands). We analysed genetic variation to estimate structure, gene flow, genetic diversity, effective size and recent demographic history of the populations. Additionally, 19 vultures were marked with satellite transmitters to track their migration routes. Results Insular populations were genetically differentiated from those of the mainland. We detected immigration in the insular populations and within the continental counterpart. We found similar levels of genetic variability between the continent and the islands, and a bottleneck analysis indicated recent sharp population declines in both archipelagos but not on the continent. Main conclusions Our study provides evidence that, in spite of significant differentiation, insular populations of highly mobile species may remain connected with the mainland. Conservation programmes should take into account population connectivity and integrate differentiated units of management within complex units of conservation that can best maintain processes and potential for evolutionary change.     

Sex-biased initial eggs favours sons in the slightly size-dimorphic Scops owl (Otus scops)

There is considerable controversy concerning the origin of Iberian populations of the Mediterranean chameleon, Chamaeleo chamaeleon . Current opinion dictates that Spanish populations result from introductions during the 18^th and 19^th centuries, with subsequent translocations from the original populations to other parts of Spain. The Portugese population in the Algarve is believed to have been introduced from Africa or Spain during the 1920s. However, Holocene remains of chameleons suggest that the Malaga population at least could have a much older origin. Analysis of sequences from the mitochondrial 16S ribosomal RNA gene of samples from the Iberian Peninsula and North Africa revealed a double origin for the Iberian population. The Mediterranean Iberian (Malaga) population is closely related to Mediterranean North African populations, with Atlantic Iberian populations more closely related to populations of the Atlantic coast of North Africa. The overall genetic differentiation and diversity observed was very low, preventing precise dating of the colonization events. However this low level of differentiation is not consistent with Plio-Pleistocene colonization, the assumed timing for a natural colonization event and suggests that chameleons were probably introduced twice by man in the recent past. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 1–7.     

Biogeographical connections between the Maghreb and the Mediterranean peninsulas of southern Europe

The glacial–interglacial cycles have caused severe range modifications of species' distributions. In Europe, thermophilic species had to retreat into geographically distinct southern refugia during glaciations. This process produced strong genetic imprints, which are still detectable by the present pattern of genetic differentiation and the distribution of regional diversity. To reveal the biogeographical imprints in the western Mediterranean, we analysed 26 populations of the butterfly Maniola jurtina spread over large areas of its European and North African distribution range. The samples were analysed using allozyme electrophoresis. We detected three genetic groups, divided into Western Europe, Central/Eastern Europe, and Italy with the Maghreb. The North African samples randomly cluster within the Italian samples. Even the population sampled in Morocco is genetically closely related to these samples and not to the geographically neighbouring Iberian ones. Parameters of genetic diversity showed similar values over the whole study area. The observed genetic pattern reflects possible glacial refugia in Europe located in the Iberian Peninsula and the Balkans. For North Africa and Italy, our data reveal a colonization of Africa originating from Italy.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 693–703.     

Phylogenetics and phylogeography of the monocot genus Baldellia (Alismataceae): Mediterranean refugia, suture zones and implications for conservation

Aquatic plants, and especially the emblematic genus Baldellia (Alismataceae), are among the most threatened organisms, due to unprecedented human-driven habitat destructions. Therefore protection plans are crucially needed and call for thoroughly documenting the genetic diversity and clarifying the taxonomy of this endangered genus. Our sampling included 282 individuals from 42 natural populations and covered the whole geographical range of the genus, across Europe and the Mediterranean. We combined sequencing of nuclear internal transcribed spacer (ITS) and chloroplastic trnL-ndhF regions with amplified fragment length polymorphism (AFLP) genotyping to investigate the Alismataceae phylogeny, and produce a phylogeography of Baldellia. Our phylogeny strongly supported the monophyly of Baldellia and placed it as the sister clade to Luronium and Alisma, therefore excluding, as previously supposed, a close genetic relatedness to the predominantly neotropical genus Echinodorus. The phylogeography of Baldellia outlined patterns consistent with a hypothesis considering glacial refugia located in the Iberian Peninsula and the Italy/Balkan region from which two distinct genetic lineages re-colonized Europe. These two lineages corresponded respectively to Baldellia ranunculoides (Italy/Balkan derived populations) and Baldellia repens (populations recovered from the Iberian Peninsula refuge), therefore supporting differences outlined between the two taxa in previous ecological and morphological studies. These results allowed clarifying taxonomic uncertainties by confirming the genetic distinctness of B. repens according to B. ranunculoides. A third lineage, Baldellia alpestris, originated and remained endemic to the mountainous regions of the Iberian Peninsula. Unexpectedly, B. repens populations collected in northern Africa, appeared to be genetically distinct from their European counterparts, this calls for further investigation to fully address their genetic and conservation status. Finally, we detected a large hybridization zone in northwestern Europe between B. repens and B. ranunculoides. These results were discussed in light of conservation approaches for Baldellia populations.     

Size‐related and diel variations in microhabitat use of three endangered small fishes in a Mediterranean coastal stream

This study analysed the microhabitat use of three endangered fish species, Andalusian toothcarp Aphanius baeticus , Iberian loach Cobitis paludica and sand smelt Atherina boyeri , in a coastal stream stretch. Plastic minnow traps were set both during daytime and at night on the bottom and at the surface. Depth and presence of effective refuge were recorded for each trap. To assess size‐related changes in microhabitat use individuals of each species were classified in three size classes. The three species preferentially used bottom positions in the water column, though this behaviour was more evident in the case of Iberian loach. While large Iberian loach remained active at night Andalusian toothcarp and sand smelt were strongly diurnal, especially larger individuals. The three species showed a clear ontogenetic change in microhabitat preferences towards deeper waters. Small Andalusian toothcarp and medium‐sized Iberian loach used deeper microhabitat in the presence of refuge. Large Andalusian toothcarp consistently preferred exposed microhabitat at any time. Andalusian toothcarp using refuge were smaller at any time, while Iberian loach followed this pattern only at night. The differential vulnerability of these species to different predators (aerial and aquatic; diurnal and nocturnal) could explain the observed patterns in microhabitat use. Fish tended to co‐occur in microhabitats either due to habitat characteristics independently of species or due to species independently of habitat. Andalusian toothcarp segregated spatio‐temporally from sand smelt and Iberian loach, but these species occurred independently of each other. According to these co‐occurrence patterns, Andalusian toothcarp would be more sensitive than Iberian loach or sand smelt to interspecific interactions.     

Reproductive strategy and ploidy determine the genetic variability of Sorbus aria

Sorbus aria (L.) Crantz (common whitebeam) from the Canary Islands has not been characterised genetically. We analysed the genetic variability of 184 individuals belonging to seven natural populations of S. aria from the Canarian Archipelago and the Iberian Peninsula. Our main aims were to obtain essential information to enable the exploration of the genetic relationship between populations from the Canary Islands and the Iberian Peninsula; to establish the existence of a spatial genetic structure and formulate appropriate management and conservation genetics strategies. Genetic variation was analysed using nine polymorphic microsatellite loci. The Canary Island populations (triploids) were found to have very low genetic variability and to be considerably differentiated from the populations from the peninsula (diploid and triploid), although with a connection to the Sierra Nevada population in the south of the Peninsula. This population, in turn, had many different genotypes, which is indicative of the existence of various origins. The level of genetic diversity was higher in all-diploid populations, which, in addition, presented a greater interpopulation gene flow, possibly the result of a prevalence of sexual reproduction. On the other hand, the triploid populations presented lower levels of genetic variability, with a significant degree of fixed heterozygosity, possibly due to asexual reproduction, mainly by apomixis. The reproductive biology and ploidy appear to be responsible for the levels of genetic variability in S. aria.     

Genome-wide comparison of African-ancestry populations from CARe and other cohorts reveals signals of natural selection

The study of recent natural selection in human populations has important applications to human history and medicine. Positive natural selection drives the increase in beneficial alleles and plays a role in explaining diversity across human populations. By discovering traits subject to positive selection, we can better understand the population level response to environmental pressures including infectious disease. Our study examines unusual population differentiation between three large data sets to detect natural selection. The populations examined, African Americans, Nigerians, and Gambians, are genetically close to one another (F_ST < 0.01 for all pairs), allowing us to detect selection even with moderate changes in allele frequency. We also develop a tree-based method to pinpoint the population in which selection occurred, incorporating information across populations. Our genome-wide significant results corroborate loci previously reported to be under selection in Africans including HBB and CD36. At the HLA locus on chromosome 6, results suggest the existence of multiple, independent targets of population-specific selective pressure. In addition, we report a genome-wide significant (p = 1.36 × 10⁻¹¹) signal of selection in the prostate stem cell antigen (PSCA) gene. The most significantly differentiated marker in our analysis, rs2920283, is highly differentiated in both Africa and East Asia and has prior genome-wide significant associations to bladder and gastric cancers.     

Invasion history of Caprella scaura Templeton, 1836 (Amphipoda: Caprellidae) in the Iberian Peninsula: multiple introductions revealed by mitochondrial sequence data

The caprellid Caprella scaura, native to the western Indian Ocean, is one striking example of a successful invader. It was first recorded in the Iberian Peninsula in Gerona (north-eastern coast of Spain), in 2005, and has recently been reported throughout the Mediterranean and Atlantic coasts of this region. The most likely vector of introduction and distribution is within the hull-fouling community on recreational craft. However, beyond the dates of detection, the introduction history of C. scaura remains unclear. Direct sequencing of mitochondrial DNA (cytochrome c oxidase subunit I) was used to compare genetic composition in native and introduced populations in order to infer the invasion history of this species. In addition, 18S rDNA sequences were used to resolve phylogenetic relationships within this species and with the morphologically closest species Caprella californica and Caprella scauroides. The high genetic divergence and population subdivision found between non-native Iberian populations together with a high level of genetic diversity in some populations indicate multiple geographical sources and introduction points for this caprellid. Our data suggest that Iberian populations may derive from at least two sources: (1) Pacific Australian, and (2) Indian Ocean, either directly, or, more likely, through stepping-stone events from central Mediterranean population(s). Atlantic Iberian populations seem to be the most recently established populations. On the other hand, this is the first study providing molecular evidence confirming C. scaura, C. californica and C. scauroides as distinct species. It also provides strong molecular evidence that C. scaura typica and C. scaura scaura correspond to the same subspecies, and the Japanese C. scaura diceros and the Chilean C. scaura spinirostris could merit specific rank.     

The molecular systematics and biogeography of the European cobitids based on mitochondrial DNA sequences

Phylogenetic inference regarding the biogeography and evolution of the family Cobitidae depends in large part on the correct interpretation of transitions between the morphological states of secondary sexual characters (e.g., the scale of Canestrini or lamina circularis). Here, we use the complete mitochondrial ATP synthase 8 and 6 and cytochrome b genes to provide an independent assessment of systematics and biogeographic relationships of species in the genus Cobitis, including geographic and subgeneric sampling of species with Canestrini's scale present, duplicated, or absent. The mtDNA-based phylogeny for the genus Cobitis provides the first formal hypothesis for the group and permits a phylogenetic-based assessment of the morphological transitions demonstrated by Canestrini's scale. Our data confirm the monophyly of the genus Cobitis and indicate that European Cobitis comprise six evolutionarily independent lineages. These lineages were defined by nucleotide synapomorphies permitting bootstrapped confidence estimates of 95% or greater and mtDNA genetic distances greater than 4.5% and correspond with moderate fidelity to the Cobitis groups defined by Bacescu (1962, Rev. Roum. Biol. 4, 435-448). The Caucasian lineage, C. cf. sibirica, represents the basal sister species of the genus Cobitis, supporting an eastern Asiatic origin of the European Cobitis: Cobitis sensu stricto, Acanestrinia, Bicanestrinia, Iberocobitis, and Cobitis calderoni. Phylogenetic relationships among Cobitis subgenera and species indicate that the ancestral condition of one scale of Canestrini was duplicated once at the origin of the Bicanestrinia lineage and has been independently lost by C. calderoni and C. elongata. The absence of the scale of Canestrini is the synapomorphy defining the subgenus Acanestrinia, but the mtDNA phylogeny indicates that Acanestrinia is not a natural group and places C. calderoni as the sister lineage to the subgenus Iberocobitis, a finding that is also geographically parsimonius.     

GENETIC DIVERGENCE IN PERIPHERALLY ISOLATED POPULATIONS OF CHAFFINCHES IN THE ATLANTIC ISLANDS

Peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Canaries, Madeira, and Azores were compared genetically with their putative ancestral stock in Iberia and Morocco, and with a population of blue chaffinches (F. teydea) from Tenerife, using protein electrophoresis of 42 loci. The continental populations are only weakly differentiated genetically (F_ST = 0.092), despite distinctive subspecific differences in plumage and morphometrics between Iberia and Morocco populations. Estimated levels of gene flow among continental populations are high enough to account for their relative genetic homogeneity, and it is unlikely that homogenizing selection is operating to mimic the effects of gene flow. In contrast, the Atlantic island populations are well differentiated genetically (F_ST = 0.321), and have diverged considerably from their continental conspecifics. The development of significant genetic differentiation within the Canaries but not the Azores likely results from smaller population sizes, very restricted gene flow, and enhanced random drift in the former populations. There is no convincing evidence in support of stronger directional selection acting on genotypes or phenotypes to reduce within-population variability in Canaries populations as proposed by Grant (1979), although other tenets of his model of island evolution are supported by our analysis. Although genetic variability is reduced in four of the Canaries populations, only the Hierro population appears to have encountered a severe bottleneck. Yet it has not differentiated markedly from the La Palma population to which it is subspecifically allied. We conclude that gradual divergence in isolated populations of small to moderate size is the most plausible explanation for the evolution of intraspecific and interspecific diversity in Atlantic island chaffinches.     

A striking example of the founder effect in the mollusc Littorina saxatilis

Two South African populations of Littorina saxatilis were examined by starch-gel electrophoresis at 16 enzyme loci and compared with 13 populations of North Atlantic saxatilis from both American and European coasts, and with six British populations of the closely related species Littorina arcana. The South African animals showed a severely reduced heterozygosity (– 0.052) compared with Atlantic populations of saxatilis ( = 0.181), and the mean genetic distance between the two areas was high ( = 0.203) compared with distances within the North Atlantic saxatilis populations (D = 0.034). In fact, the saxatilis from South Africa were genetically more distant from the North Atlantic samples of L. saxatilis than were the arcana from British shores. The reduced genetic heterozygosity and genetic divergence of the South African populations is attributed to founder effects following a postulated recent introduction by man.