The genetic population structure of Buthus occitanus (Scorpiones: Buthidae) across the Strait of Gibraltar: calibrating a molecular clock using nuclear allozyme variation

I assess here the importance of the Strait of Gibraltar as a barrier to gene flow for populations of the scorpion Buthus occitanus . This polytypic buthid scorpion occurs in Europe and in North Africa where it is morphologically more diverse. The phylogenetic relationship between B. occitanus populations across the Strait of Gibraltar is investigated by nuclear allozymes analysis (15 loci scored). Phylogenetic analysis based on estimated gene frequency data results in a tree topology that divides the populations into three clades, i.e. a European, an Atlas (= Morocco samples) and a Tell-Atlas clade (= Tunisian samples). The Tell-Atlas clade grouped with the European clade with a rather high bootstrap support of 70%. Within these clades low levels of genetic variability are observed. Calibrating a molecular clock under the assumption that the European populations are autochthonous and have been isolated from the North African for at least 5.33 Myr reveals a divergence rate of 0.060 genetic distance (D) per Myr estimated between European and Moroccan samples and 0.036D Myr⁻¹ between European and Tunisian samples, respectively.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 519–534.     

Molecular differentiation between coexisting species of Taraxacum sect. Erythrosperma (Asteraceae) from populations in south-east and west Germany

The present study focuses on the genetic differentiation between six coexisting species of Taraxacum sect. Erythrosperma . Twenty-six individuals of T. rubicundum , T. brachyglossum , T. tortilobum , T. lacistophyllum , T. parnassicum and T. scanicum were sampled at 14 locations in south-east (Bavaria) and west Germany (Eifel). Using random amplified polymorphic DNA (RAPD) analysis 139 fragments were amplified. Nineteen of these were found to be species specific. The level of polymorphic bands within the different Taraxacum species varied between 0% and 15.8%. An analysis of molecular variance (AMOVA) revealed strong differentiation between the investigated species: 14.3% of the molecular variance were found within the species, while 85.7% of the genetic variability was observed between the species. However, a geographical differentiation between the different regions could not be observed. All analysed species were clearly separated from each other in a cluster analysis using the neighbour joining technique. The observed strong genetic differentiation between the Taraxacum species clearly supports the agamospecies concept of Taraxacum sect. Erythrosperma , which is at present mainly based on morphological and cytological criteria  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 109–117.     

Genetic divergence in the superspecies Manacus

The bearded manakins in the genus Manacus are lekking, neotropical passerines. Male plumage colour varies with geographical location and classification is based solely on these plumage patterns. It has recently been suggested that in this group of birds, plumage patterns may be a misleading taxonomic character. In this study we used microsatellite variation in a collection of museum samples to establish the amount of genetic divergence between the previously described bearded manakin species/subspecies. We found substantial genetic substructuring between species/subspecies and that plumage patterns indeed may be a misleading taxonomic character because the presence of yellow in male nuptial plumage is found in most divergent forms. We did not detect a significant isolation by distance relationship although the P -value was close to significance. Physical barriers such as rivers and mountains may affect gene flow and play a role in shaping genetic structure of the genus Manacus . Accordingly, boundaries between species/subspecies often coincide with large rivers, mountains and seas.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 439–447.     

Implications of contrasting patterns of genetic variability in two vespertilionid bats from the Indonesian archipelago

Wallacean island populations of two Vespertilioninae bats, Myotis muricola and Scotophilus kuhlii , which have similar geographical ranges, showed marked contrast in the amount and pattern of genetic variation. Within islands, genetic variation was on average much higher in M. muricola but declined from west to east, whereas all populations of S. kuhlii had uniformly low levels of genetic variation by mammalian standards. S. kuhlii showed little genetic differentiation between islands and estimates of gene flow were substantial whereas island populations of M. muricola differed markedly and there was a strong isolation-by-distance effect associated with the extent of the sea crossing between islands. Furthermore, the lower mean heterozygosity and small genetic distances between eastern island populations of M. muricola is evidence that there has been a bottleneck associated with the colonization of this area. The attenuation of genetic diversity to the east is also seen in some other mammalian species and may indicate limits to dispersal and have implications for species management. The patterns of variability in S. kuhlii may be a consequence of its strong dispersal capacity and close association with human activity, which, together with other factors, suggest a panmictic population.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 421–431.     

Stability of genetic structure and effective population size inferred from temporal changes of microsatellite DNA polymorphisms in the land snail Helix aspersa (Gastropoda: Helicidae)

Temporal evolution of genetic variability may have far-reaching consequences for a diverse array of evolutionary processes. Within the polders of the Bay of Mont-Saint-Michel (France), populations of the land snail Helix aspersa are characterized by a metapopulation structure with occasional extinction processes resulting from farming practices. A temporal survey of genetic structure in H . aspersa was carried out using variability at four microsatellite loci, in ten populations sampled two years apart. Levels of within-population genetic variation, as measured by allelic richness, H _e or F _is , did not change over time and similar levels of population differentiation were demonstrated for both sampling years. The extent of genetic differentiation between temporal samples of the same population established (i) a stable structure for six populations, and (ii) substantial genetic changes for four populations. Using classical F -statistics and a maximum likelihood method, estimates of the effective population size ( N _e) illustrated a mixture of stable populations with high N _e, and unstable populations characterized by very small N _e estimates (of 5–11 individuals). Owing to human disturbances, intermittent gene flow and genetic drift are likely to be the predominant evolutionary processes shaping the observed genetic structure. However, the practice of multiple matings and sperm storage is likely to provide a reservoir of variability, minimizing the eroding genetic effects of population size reduction and increasing the effective population size.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 89–102.     

Cryptic species within the freshwater isopod Mesamphisopus capensis (Phreatoicidea: Amphisopodidae) in the Western Cape, South Africa: allozyme and 12S rRNA sequence data and morphometric evidence

The freshwater phreatoicidean isopod Mesamphisopus capensis has been regarded as the most widespread of the four Mesamphisopus species occurring in the Western Cape, South Africa. To determine whether this species was monotypic across its distribution over two mountainous regions, separated by a low-lying coastal plain remnant, genetic differentiation among populations from 11 localities was studied through allozyme electrophoresis of 12 loci and sequencing of a 338-bp 12S rRNA mtDNA fragment from representative individuals. Populations of the two regions were separated by a mean identity value of 0.477. Fixed allele differences at two loci distinguished these regions. Estimates of θ indicated substantial differentiation among populations across the entire sample, as well as within each of the regions. Topologies derived through parsimony and neighbour joining supported the monophyly of the two regions. On the basis of these topologies, allele frequencies and an allozyme dendrogram, five groups were identified. Discriminant function analyses, performed on body and pereopod variables independently, revealed these groups to be well differentiated with a high rate of correct a posteriori reclassification. Using genetic distance criteria these five distinct forms may be considered to be putative species. From a conservation perspective, the two regions can be seen to represent two evolutionarily significant units, while the five groups should be regarded as management units.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 235–253.     

A preliminary palynological characterization of Spanish thyme honeys

Pollen was analysed in 25 thyme honey samples from Spain. The honey samples were provided by professional beekeepers, and pollen was obtained by centrifugation. A minimum of 8% of Thymus sp. pollen is considered sufficient to typify a honey as unifloral when considering pollen grains from anemophilous and nectarless plants. The quantitative analysis showed that nectar is the main honey source in the samples studied. The qualitative analysis of the samples revealed the presence of 53 taxa belonging to 27 families. The Spanish thyme honeys of the studied region are characterized by their medium to high numbers of grains of pollen (NGP; mean = 155 000) and their medium to high content of honeydew indicator elements (HDE; mean = 8300). Genista type (present in 100% of the samples), Echium plantagineum and Reseda luteola (90%), Eucalyptus type (80%), Calendula type and Olea europaea (75%), and Cistus monspeliensis (70%) may be mentioned among the characteristic accompanying species of this honey type.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 323–330.     

Allozymic differentiation of Tunisian populations of Androctonus species and Buthus occitanus (Scorpiones: Buthidae)

We examined the population structure and phylogeny of Buthus occitanus (Amoreux, 1789), Androctonus amoreuxi (Audouin & Savigny 1812) and Androctonus australis (Linnaeus, 1758) in Tunisia, using horizontal starch gel and polyacrylamide gel electrophoresis to resolve allozymes at 18 loci. Populations of B. occitanus and A. amoreuxi showed little genetic variation, whereas A. australis populations showed moderate genetic variation. Moreover, despite the restricted dispersal abilities of these species, the allozyme analysis indicated very low levels of intraspecific population differentiation. The Island model of population differentiation best fitted the intraspecific variation, both for mainland populations and for the slightly more differentiated populations from two Mediterranean islands. No diagnostic, fixed alleles were observed between the species of Androctonus , which concurs with the results of nuclear ribosomal DNA analysis. Two subspecies of A. australis , A. a. garzonii Goyffon & Lamy, 1973 and A. a. hector Vachon, 1948, were not resolved by a phylogenetic analysis of the allozyme data. The most likely reasons for the slight population differentiation are either low mutation rates or recent range extensions promoted by human activities. The possibility of indirect gene flow between Androctonus species should also be explored.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 255–265.     

Current and historic gene flow of the sand goby Pomatoschistus minutus on the European Continental Shelf and in the Mediterranean Sea

Phylogeographical patterns of the sand goby Pomatoschistus minutus (Gobiidae, Teleostei) were studied by means of sequence and single-stranded conformational polymorphism analysis of a 283-bp fragment of the cytochrome b locus of the mtDNA. A total of 228 individuals sampled at 13 sites throughout the species's distributional range revealed a moderate level of diversity and a low but significant level of overall genetic differentiation at all but one site. The goby sample from the Adriatic Sea differed in sequence by approximately 10% from the Atlantic P. minutus and is thought to belong to a cryptic species of the genus Pomatoschistus . Limited genetic differentiation with a weak pattern of isolation-by-distance was recorded throughout the distributional range of the typical P. minutus . Phylogeographical analysis suggested a contiguous range expansion in the Atlantic and Baltic basins during the Eemian and evidence for a glacial refugium in the southern North Sea during the Weichselian. In P. minutus from the western Mediterranean Sea a high number of endemic haplotypes as well as the most common Atlantic haplotype were recorded in appreciable frequencies. This might be explained by secondary contact between different mitochondrial lineages, which evolved in allopatry.   © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 561–576.     

Molecular taxonomy and phylogeography of Miniopterus schreibersii (Kuhl, 1817) (Chiroptera: Vespertilionidae), in the Eurasian transition

Miniopterus schreibersii is a polytypic bat species, with one of the widest distribution ranges among the mammals. We studied the genetic differentiation and taxonomy of this species in the transition zone between south-eastern Europe and Anatolia (in Asia), where two subspecies have been described. The results indicated a sharp genetic break between the samples from western Anatolia and south-eastern Europe and those of eastern Anatolia. In addition, the samples from western Anatolia and south-eastern Europe were seen to be reciprocally monophyletic, although the differentiation was less drastic. These patterns of genetic differentiation suggest the presence of two distinct groups within the M. schreibersii complex in the region, concordant with previous subspecific recognition. The cause of this genetic break is most likely differentiation in separate glacial refugia followed by secondary contact. However, more samples are needed to assess whether these represent different species, as well as to understand more clearly the causes of this differentiation.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 577–582.     

Subtle population genetic structure in the Hawaiian grouper, Epinephelus quernus (Serranidae) as revealed by mitochondrial DNA analyses

The endemic Hawaiian grouper, Epinephelus quernus , is a commercially important species experiencing intense fishing pressure in part of its distributional range. We examined population genetic structure with 398 base pairs of the mitochondrial control region across a large portion of the range of E. quernus , spanning approximately 2000 km of the Hawaiian archipelago. Examination of genetic diversity shows that Gardner Island, situated midway along the island chain, harbours the most diverse haplotypes. F -statistics and Bayesian estimates of migration also reveal the mid-archipelago as genetically differentiated, where the first significant break among adjacent pairs of populations lies between the islands of Nihoa and Necker. Most island comparisons beyond Necker and Gardner to the north-west and among the lower five islands to the south-east show little to no genetic differences. Evidence of historical population expansion across the islands was also found by Maximum Likelihood analyses. The results suggest that management should be structured to reflect the genetic differentiation and diversity in the mid-archipelago, the patterns of which may be associated with oceanic current patterns.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 449–468.     

Population genetic structure of New Zealand's endemic corophiid amphipods: evidence for allopatric speciation

Allozyme electrophoresis was used to examine population genetic structure at inter- and intraspecific levels for the New Zealand endemic corophiid amphipods, Paracorophium lucasi and P. excavatum. Individuals were collected from estuarine and freshwater habitats from North, South and Chatham Islands. Analyses of genetic structure among interspecific populations indicated clear allelic differentiation between the two Paracorophium species (Nei's genetic distance, D  = 1.62), as well as considerable intraspecific substructuring ( D  = 0.15–0.65). These levels of divergence are similar to interspecific levels for other amphipods and it is proposed that at least two groups from the P. lucasi complex and three from the P. excavatum complex correspond to sibling species. In most cases allopatry can account for the differentiation among the putative sibling species. For populations that share a common coastline we found low levels of differentiation and little or no correlation with geographical distance, suggesting that gene flow is adequate to maintain homogeneous population genetic structure. By contrast, populations on separate coastlines (i.e. isolated by land) showed moderate levels of geographical differentiation indicating restricted gene flow. The juxtaposition of population genetic and biogeographical data for Paracorophium in conjunction with the geological record infers past histories of glacial extirpation, and possible isolating effects of sea-level and landmass changes that have occurred throughout the Plio-Pleistocene.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 119–133.     

Polyploidy and the sexual system: what can we learn from Mercurialis annua?

The evolutionary success of polyploidy most directly requires the ability of polyploid individuals to reproduce and transmit their genes to subsequent generations. As a result, the sexual system (i.e. the mating system and the sex allocation of a species) will necessarily play a key role in determining the fate of a new polyploid lineage. The effects of the sexual system on the evolution of polyploidy are complex and interactive. They include both aspects of the genetic system, the genetic load maintained in a population and the ecological context in which selection takes place. Here, we explore these complexities and review the empirical evidence for several potentially important genetic and ecological interactions between ploidy and the sexual system in plants. We place particular emphasis on work in our laboratory on the European annual plant Mercurialis annua , which offers promising scope for detailed investigations on this topic. M. annua forms a polyploid complex that varies in its sexual system from dioecy (separate sexes) through androdioecy (males and hermaphrodites) to functional hermaphroditism.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 547–560.     

The lichenicolous Opegrapha species (Roccellaceae, Ascomycota) with 3-septate ascospores on Pertusaria and Ochrolechia

The Opegrapha species with 3-septate ascospores growing on Pertusaria and Ochrolechia are revised. Two species are recognized: Opegrapha anomea (of which O. pertusariae , O. quaternella , O. wetmorei and possibly Leciographa weissii are considered to be synonyms), and O. blakii Ertz & Diederich sp. nov. described from a sterile lichen with an Ochrolechia -like thallus, known from Ecuador and Venezuela. Opegrapha anomea and several related lichenicolous species with roundish or irregular, often multilocular ascomata are morphologically intermediate between Opegrapha and Plectocarpon , and might represent a distinct genus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144, 235−241.     

Geographic patterns of genetic variation in four Neotropical rodents: conservation implications for small game mammals in French Guiana

The Neotropics contributes significantly to global biodiversity yet little is known about its mammalian fauna. Given our limited knowledge, concerns are that extinctions at the local level may cause a decline in genetic diversity through the loss of unique alleles. Here we report the geographical population structure of four rodent species from French Guiana: Dasyprocta leporina , Agouti paca , Proechimys cayennensis and P. cuvieri . Two of them ( Dasyprocta and Agouti ) are commonly harvested throughout the country for subsistence and/or game hunting. Cytochrome b and control region sequences revealed the presence of a number of maternal lineages occurring in sympatry throughout French Guiana. We applied two cytochrome b rodent clock calibrations (7.5% and 12% per million years) to our data, and placed the divergence of the maternal lineages to between 160 000 and 260 000 years ago for the more diverse D. leporina and P. cuvieri , and between 66 000 and 46 000 years ago for the more recent A. paca and P. cayennensis . The clades appear to be widespread throughout South America as indicated by specimens sampled in Brazil, Peru, and Venezuela. At the population level, AMOVA revealed little or no geographical structure within French Guiana. However, this conclusion is based on a single genetic marker and relatively few specimens.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 203–218.     

Genetic differentiation on multiple spatial scales in an ecotype-forming marine snail with limited dispersal: Littorina saxatilis

The population genetic structure of marine species lacking free-swimming larvae is expected to be strongly affected by random genetic drift among populations, resulting in genetic isolation by geographical distance. At the same time, ecological separation over microhabitats followed by direct selection on those parts of the genome that affect adaptation might also be strong. Here, we address the question of how the relative importance of stochastic vs. selective structuring forces varies at different geographical scales. We use microsatellite DNA and allozyme data from samples of the marine rocky shore snail Littorina saxatilis over distance scales ranging from metres to 1000 km, and we show that genetic drift is the most important structuring evolutionary force at distances > 1 km. On smaller geographical scales (< 1 km), divergent selection between contrasting habitats affects population genetic structure by impeding gene flow over microhabitat borders (microsatellite structure), or by directly favouring specific alleles of selected loci (allozyme structure). The results suggest that evolutionary drivers of population genetic structure cannot a priori be assumed to be equally important at different geographical scales. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 31–40.     

Phylogeography of the freshwater crayfish Cherax destructor Clark (Parastacidae) in inland Australia: historical fragmentation and recent range expansion

The yabby, Cherax destructor Clark, is the most widespread species in the most widespread genus of Australian freshwater crayfish. It has a distribution that spans several distinct drainage basins and biogeographical regions within semiarid and arid inland Australia. Here we report a study designed to investigate patterns of genetic variation within the species and hypotheses put forward to account for its extensive distribution using DNA sequences from the mitochondrial 16S rRNA gene region. Results of phylogenetic analyses contradicted previous allozyme data and revealed relatively deep phylogenetic structure in the form of three geographically correlated clades. The degree of genetic divergences between clades (8–15 bp) contrasted with the relatively limited haplotype diversity within clades (1–3 bp). Network-based analyses confirmed these results and revealed genetic structure on both larger and more restricted geographical scales. Nevertheless some haplotypes and 1-step clades had large distributions, some of which crossed boundaries between river basins and aquatic biogeographical regions. Thus both older and more recent historical processes, including fragmentation on a larger geographical scale and more recent range expansion on a local scale, appear to be responsible for the observed pattern of genetic variation within C. destructor . These results support elements of alternative hypotheses previously put forward to account for the evolutionary history of C. destructor and the origin of its large distribution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 539–550.     

Growth pattern of Carex arenaria (Cyperaceae)

The origin of the orthotropic aerial shoot from the plagiotropic, horizontal rhizome of Carex arenaria L. (Cyperaceae) was studied. A previously proposed adnation or fusion of the first internode of the orthotropic shoot and the fourth internode of the plagiotropic shoot could not be anatomically confirmed. The rhizome branches sympodially and the point of branching results from intercalary meristematic activity in the plagiotropic shoot.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 145–151.