Life history divergence of sympatric diploid and polyploid populations of brine shrimp Artemia parthenogenetica

In order to study how polyploidy affects life history patterns in animals, we have examined sympatric diploid and polyploid brine shrimp (Artemia parthenogenetica) from China, Italy and Spain under laboratory conditions. At optimal temperature and salinity (25°C and 90 ppt), diploids from the three populations had much higher intrinsic rates of increase, higher fecundity, faster developmental rates, and larger brood sizes than their sympatric polyploids. The Chinese and Italian populations were selected for further analysis to determine the life history responses of diploids and polyploids to temperature and salinity changes. Under intermediate and high salinities, Chinese and Italian polyploids produced most of their offspring as dormant cysts while their sympatric diploids produced most of their offspring as nauplii. This relationship is reversed in the Spanish diploid-polyploid complex. For the Chinese population at 25° C, pentaploid clones had higher developmental rates than diploid clones at 35 ppt; at 90 ppt, diploid clones had higher developmental rates than the pentaploids. Italian diploids and tetraploids had different responses to variation in both temperature (25° C and 31° C) and salinity (30 ppt and 180 ppt). Our results demonstrate that relative fitness of the two cytotypes is a function of environmental conditions and that sympatric diploids and polyploids respond differently to environmental changes. Chinese and Italian polyploids are expected to have lower fitness than their sympatric diploids when the physical environment is not stressful and when intraspecific competition is important. However, polyploids may have advantages over sympatric diploids in stressful habitats or when they encounter short-term lethal temperatures. These results suggest that polyploid Artemia have evolved a suite of life-history characteristics adapting them to environments that contrast to those of their sympatric diploids.     

RECONSTRUCTING RETICULATION HISTORY IN A PHYLOGENETIC FRAMEWORK AND THE POTENTIAL OF ALLOPATRIC SPECIATION DRIVEN BY POLYPLOIDY IN AN AGAMIC COMPLEX IN CRATAEGUS (ROSACEAE)

Polyploidy plays a prominent role in the speciation process in plants. Many species are known to be part of agamic complexes comprising sexual diploids and more or less exclusively asexual polyploids. However, polyploid formation has been studied in very few cases, primarily because of the challenges in examining these cases phylogenetically. In this study, we demonstrate the use of a variety of phylogenetic approaches to unravel origins and infer reticulation history in a diploid–polyploid complex of black‐fruited Crataegus. The tree approaches are shown to be useful in testing alternative hypotheses and in revealing genealogies of nuclear genes, particularly in polyploid organisms that may contain multiple copies. Compared to trees, network approaches provide a better indication of reticulate relationships among recently diverged taxa. Taken together, our data point to both the autopolyploid and allopolyploid origins of triploids in natural populations of Crataegus suksdorfii, whereas tetraploids are formed via a triploid bridge, involving the backcross of allotriploid offspring with their diploid C. suksdorfii parent, followed by gene introgression from sympatric C. douglasii. Our findings provide empirical evidence for different pathways of polyploid formation that are all likely to occur within natural populations and the allopatric establishment of neopolyploids subsequent to their formation.     

Genetic diversity in sexual diploid and apomictic tetraploid populations of Paspalum notatum situated in sympatry or allopatry

Paspalum notatum is a subtropical grass widely distributed in the temperate areas of America. Diploids are sexual while polyploids give rise to clonal seeds through aposporous apomixis. RAPD markers were used to analyze the genetic structure of three natural populations: i) diploids reproducing sexually (R2X); ii) sympatric apomictic tetraploids collected in the vicinity of the diploids (R4X); iii) allopatric apomictic tetraploids growing in isolation (C4X). The apomictic reproduction rate was evaluated by the use of molecular markers in progeny tests, while chromosome-counting allowed the verification of ploidy levels. Data revealed that the R4X group presented a variation considerably higher than that observed for C4X. Jaccards coefficients were used to produce a cluster diagram using the UPGMA method. All but one tetraploid genotypes grouped together and were associated to diploid genotype A21. The possibility of occasional generation of novel tetraploid clones from the interaction between tetraploid and diploid individuals is discussed.     

Clonal diversity in the agamospermous polyploids ofTaraxacum hondoense in northern Honshu,Japan

Chromosome numbers and allozyme variations were surveyed in 74 polyploid populations ofTaraxacum hondoense, in northern Honshu, Japan. Most of the populations (94.4%) consisted of triploid (2n=24), indicating the predominance of this ploidy level. Approximately 42.6% were found to contain tetraploid (2n=32), and a few plants were pentaploid (2n=40). Electrophoretic analysis at6 Pgdh-1 revealed twelve phenotypes with four alleles (including one putative null allele). The triploids showed excessive heterozygosity (82.4%) and all of the tetraploids and pentaploids were heterozygote. Phenotype IV was the most frequent and widely distributed in northern Honshu. Forty five percent of the populations were found to contain multiple phenotypes at 6Pgdh-1. A total of 21 clones were distinguished using three polymorphic loci (6Pgdh-1, Got andMdh), and a considerable amount of clonal diversity was detected both within and among polyploid populations ofT. hondoense. Factors causing multiclonality in agamospermous polyploids are discussed.     

Where are the glacial refugia in Europe? Evidence from pteridophytes

In this paper we demonstrate that, by investigating polyploid complexes in Asplenium, it is possible to locate the areas in Europe that are southern glacial rcfugia, and arc likely to have been so since the beginning of the Pleistocene during the consecutive cold and warm periods in Europe. Identification and conservation of these specific areas that serve as safe havens for plants, and perhaps animals, is of paramount importance for the maintenance of European biodiversity because Man's activities arc resulting in an ever-increasing loss of natural habitats and putting diversity at risk. The genus Asplenium in Europe comprises some 50 taxa: half of these are diploid while the other half arc polyploids derived from the diploids. All aspleniums in Europe are (small) rock ferns with high substrate specificity. Today, most of mainland Europe, Scandinavia and the British Isles has been colonized by polyploid Asplenium species, while the diploids that gave rise to these polyploids are distributed around (and more or less confined to) the Mediterranean Basin. In the tetraploids genetic variation is partitioned mostly between sites, whereas diploids show a high degree of genetic variation both within and between sites. The tctraploid taxa seem capable of single spore colonization via intragametophytic selfing, but the diploid taxa appear to be predominantly outbreeding. For most diploids at least two gametophytes, produced by different spores, have to be present to achieve fertilization and subsequent sporophyte formation for the successful colonization of a new site. This results in a slower rate of colonization. The formation of auto- and allopolyploid taxa from diploid communities appears to have been a recurrent and common feature in Europe. Minority cytotypc exclusion is likely to prevent the establishment of tetraploids within the diploid communities, but spores from tetraploids can establish populations outside the diploid communities. The differences between colonization abilities of tctraploid and ancestral diploid taxa, resulting from their different breeding systems, has prevented the merging and mingling of their ranges and led to the establishment of contact/ hybrid zones. This has resulted in the restriction of diploid populations to ancient glacial rcfugia and the colonization of the rest of Europe by polyploids. Mapping the current distribution of these diploid communities and comparing the genetic diversity within and between outbreeding diploid Asplenium taxa allows us to define the area, age and historical biogcography of these rcfugia and to assess their importance for present day genetic and species diversity in Europe.     

A CYTOGEOGRAPHIC STUDY OF CHAENACTIS DOUGLASII (COMPOSITAE,HELENIEAE)

Analysis of 512 plants derived from 200 populations shows that the widely distributed western North American Chaenactis douglasii species-complex consists of diploids (n = 6), triploids, tetraploids, and hexaploids. Microsporocytes were the source of most of the chromosome counts. About 9% of all plants examined had one or more full-sized extra chromosomes. Multivalents, usually a ring or chain of four chromosomes, were almost entirely restricted to polyploids, where one or more were identified in 38% of the tetraploids and 33% of the hexaploids. With two exceptions, diploids and polyploids were not found in the same population. Frequencies of diploid, triploid, tetraploid, and hexaploid populations were, respectively, 34, 1.5, 55 and 9.5%. With significant exceptions, diploid populations predominate in the Pacific and Rocky Mountain Systems, whereas polyploid ones are most frequent in the intervening plateaus. Ploidy level is correlated with age of substrate, rather than with climate, elevation, vegetation, or soil type. Range, morphology, ploidy level, and meiotic behavior suggest that var. achilleifolia tetraploids and hexaploids are descendents of hybrids between other variants of the complex. The diploid-tetraploid-hexaploid geographic distribution and the age of the substrates where each tends to occur suggest that the complex evolved in late Cenozoic time in response to major climatic and geologic changes that induced migration and hybridization. The hybrid derivatives, stabilized by polyploidy and tolerant of increasing aridity, came to occupy newly available habitats in areas disturbed by volcanic activity and glacial or glacial-related processes.     

PLANT POLYPLOIDY AND POLLINATION: FLORAL TRAITS AND INSECT VISITS TO DIPLOID AND TETRAPLOID HEUCHERA GROSSULARIIFOLIA

In many polyploid species, polyploids often have different suites of floral traits and different flowering times than their diploid progenitor species. We hypothesized that such differences in floral traits in polyploids may subsequently affect their interactions with pollinating and other insect visitors. We measured floral morphology and flowering phenology in 14 populations of diploid and autotetraploid Heuchera grossulariifolia Rydb. (Saxifragaceae), determined if repeated evolution of independent polyploid lineages resulted in differentiation in floral morphology among those lineages, and ascertained if there was a consistent pattern of differentiation among genetically similar diploid and autotetraploid populations. In addition, we evaluated the differences in suites of floral visitors within a natural community where diploids and autotetraploids occur sympatrically. Overall, flowers of autotetraploid plants were larger and shaped differently than those of diploids, had a different flowering phenology than that of diploids, and attracted different suites of floral visitors. In comparison with flowers of diploids, tetraploid floral morphology varied widely from pronounced differences between cytotypes in some populations to similar flower shapes and sizes between ploidal levels in other populations. Observations of floral visitors to diploids and autotetraploids in a natural sympatric population demonstrated that the cytotypes had different suites of floral visitors and six of the 15 common visitors preferentially visited one ploidy more frequently. Moreover, we also found that floral morphology differed among independent autotetraploid origins, but there was no consistent pattern of differentiation between genetically similar diploid and autotetraploid populations. Hence, the results suggest that the process of polyploidization creates the potential for attraction of different suites of floral visitors. Multiple origins of polyploidy also presents the opportunity for new or different plant-insect interactions among independent polyploid lineages. These differences in turn may affect patterns of gene flow between diploids and polyploids and also among plants of independent polyploid origin. Polyploidy, therefore, may result in a geographic mosaic of interspecific interactions across a species' range, contributing to diversification in both plant and insect groups.     

A CYTOGEOGRAPHIC STUDY OF ERIOPHYLLUM LANATUM (COMPOSITAE,HELENIEAE)

Analysis of 368 plants derived from 239 natural populations showed that this taxonomically perplexing and wide-ranging species-complex consists of diploids (n = 8), tetraploids, hexaploids and octoploids. Microsporocytes were the source of most of the chromosome counts. Meiosis was basically regular. Multivalent formation was uncommon, but 11 % of all the plants examined had one or more full-sized extra chromosomes. The frequency of plants with extra chromosomes varied significantly among the taxa, from 0 (five varieties) to over 20 % (two varieties). Except in one instance, where one population yielded a diploid and a triploid, different ploidy levels were not found in the same population. The frequency of diploid, tetraploid, hexaploid and octoploid populations was, respectively, 71, 22, 4 and 2%. Variety obovatum appears to be exclusively diploid, and var. aphanactis exclusively tetraploid. Diploids and one or more polyploid levels occurred in the other taxa. No correlation was found between polyploidy and geological history, soils, topography or climate, nor were the polyploids more widely distributed than the diploids. Some of the polyploid populations seem to have been derived from inter-varietal hybridizations, but others do not. The complex has a “pillar” structure in which 10 diploid taxa support a three-level polyploid superstructure. The available evidence suggests that the major role of polyploidy here has been to stabilize the products of intra- and inter-varietal hybridizations.     

Pollinator-mediated assortative mating in mixed ploidy populations of Chamerion angustifolium (Onagraceae)

Establishment of polyploid individuals within diploid populations is theoretically unlikely unless polyploids are reproductively isolated, pre-zygotically, through assortative pollination. Here, we quantify the contribution of pollinator diversity and foraging behaviour to assortative pollen deposition in three mixed-ploidy populations of Chamerion angustifolium (Onagraceae). Diploids and tetraploids were not differentiated with respect to composition of insect visitors. However, foraging patterns of the three most common insect visitors (all bees) reinforced assortative pollination. Bees visited tetraploids disproportionately often and exhibited higher constancy on tetraploids in all three populations. In total, 73% of all bee flights were between flowers of the same ploidy (2x–2x, 4x–4x); 58% of all flights to diploids and 83% to tetraploids originated from diploid and tetraploid plants, respectively. Patterns of pollen deposition on stigmas mirrored pollinator foraging behaviour; 73% of all pollen on stigmas (70 and 75% of pollen on diploid and tetraploid stigmas, respectively) came from within-ploidy pollinations. These results indicate that pollinators contribute to high rates of pre-zygotic reproductive isolation. If patterns of fertilization track pollen deposition, pollinator–plant interactions may help explain the persistence and spread of tetraploids in mixed-ploidy populations.     

Autopolyploidy in Dactylis glomerata L.: further evidence from studies of chloroplast DNA variation

Summary Chloroplast DNA variation has been used to examine some of the maternal lineages involved in the evolution of the intraspecific polyploid complex, Dactylis glomerata L. Diploid (2x) and tetraploid (4x) individuals were collected from natural populations of the subspecies glomerata (4x), marina (4x) and lusitanica (2x), as well as from sympatric 2x/4x populations of the Galician type. Digestion of their ctDNA with 11 restriction endonucleases revealed enough variation to characterise three ctDNA variants, designated MBMK, MBmK and mBMK. The distribution of these ctDNA variants reflects different stages in their spread among the populations. The MBMK ctDNA variant predominated at both ploidy levels in subspecies glomerata, lusitanica and marina, and in recent tetraploid Galician/glomerata hybrids. The MBmK variant was detected in a single tetraploid individual and probably results from a relatively recent mutation. Fixation of the mBMK minority variant in the diploid and tetraploid Galician populations adds to the evidence concerning the possible origin of the Galician tetraploids. It means that the Galician diploids were maternal ancestors of the tetraploids. This result complements evidence from earlier studies based on morphology or biochemical markers, and reduces the likelihood that the tetraploids arose by hybridisation between an ancient Galician diploid and an alien tetraploid. It is, however, consistent with a true autopolyploid origin of the tetraploids.     

Pollen competition as a unilateral reproductive barrier between sympatric diploid and tetraploid Chamerion angustifolium

Speciation requires the evolution of barriers to gene exchange between descendant and progenitor populations. Cryptic reproductive barriers in plants arise after pollination but before fertilization as a result of pollen competition and interactions between male gametophytes and female reproductive tissues. We tested for such gametic isolation between the polyploid Chamerion angustifolium and its diploid progenitor by conducting single (diploid or tetraploid) and mixed ploidy (1 : 1 diploid and tetraploid) pollinations on both cytotypes and inferring siring success from paternity analysis and pollen-tube counts. In mixed pollinations, polyploids sired most (79%) of their own seeds as well as those of diploids (61%) (correcting for triploid block, siring success was 70% and 83%, respectively). In single donor pollinations, pollen tubes from tetraploids were more numerous than those from diploids at four different positions in each style and for both diploid and tetraploid pollen recipients. The lack of a pollen donor x recipient interaction indicates that the tetraploid siring advantage is a result of pollen competition rather than pollen-pistil interactions. Such unilateral pollen precedence results in an asymmetrical pattern of isolation, with tetraploids experiencing less gene flow than diploids. It also enhances tetraploid establishment in sympatric populations, by maximizing tetraploid success and simultaneously diminishing that of diploids through the production of inviable triploid offspring.     

Attack of the clones: reproductive interference between sexuals and asexuals in the Crepis agamic complex

Negative reproductive interactions are likely to be strongest between close relatives and may be important in limiting local coexistence. In plants, interspecific pollen flow is common between co‐occurring close relatives and may serve as the key mechanism of reproductive interference. Agamic complexes, systems in which some populations reproduce through asexual seeds (apomixis), while others reproduce sexually, provide an opportunity to examine effects of reproductive interference in limiting coexistence. Apomictic populations experience little or no reproductive interference, because apomictic ovules cannot receive pollen from nearby sexuals. Oppositely, apomicts produce some viable pollen and can exert reproductive interference on sexuals by siring hybrids. In the Crepis agamic complex, sexuals co‐occur less often with other members of the complex, but apomicts appear to freely co‐occur with one another. We identified a mixed population and conducted a crossing experiment between sexual diploid C. atribarba and apomictic polyploid C. barbigera using pollen from sexual diploids and apomictic polyploids. Seed set was high for all treatments, and as predicted, diploid–diploid crosses produced all diploid offspring. Diploid–polyploid crosses, however, produced mainly polyploidy offspring, suggesting that non‐diploid hybrids can be formed when the two taxa meet. Furthermore, a small proportion of seeds produced in open‐pollinated flowers was also polyploid, indicating that polyploid hybrids are produced under natural conditions. Our results provide evidence for asymmetric reproductive interference, with pollen from polyploid apomicts contributing to reduce the recruitment of sexual diploids in subsequent generations. Existing models suggest that these mixed sexual–asexual populations are likely to be transient, eventually leading to eradication of sexual individuals from the population.     

The association between polyploidy and clonal reproduction in diploid and tetraploid Chamerion angustifolium

Clonal reproduction is associated with the incidence of polyploidy in flowering plants. This pattern may arise through selection for increased clonality in polyploids compared to diploids to reduce mixed‐ploidy mating. Here, we test whether clonal reproduction is greater in tetraploid than diploid populations of the mixed‐ploidy plant, Chamerion angustifolium, through an analysis of the size and spatial distribution of clones in natural populations using AFLP genotyping and a comparison of root bud production in a greenhouse study. Natural tetraploid populations (N = 5) had significantly more AFLP genotypes ( = 10.8) than diploid populations ( = 6.0). Tetraploid populations tended to have fewer ramets per genotype and fewer genotypes with >1 ramet. In a spatial autocorrelation analysis, ramets within genotypes were more spatially aggregated in diploid populations than in tetraploid populations. In the greenhouse, tetraploids allocated 90.4% more dry mass to root buds than diploids, but tetraploids produced no more root buds and 44% fewer root buds per unit root mass than diploids. Our results indicate that clonal reproduction is significant in most populations, but tetraploid populations are not more clonal than diploids, nor are their clones more spatially aggregated. As a result, tetraploids may be less sheltered from mixed‐ploidy mating and diploids more exposed to inbreeding, the balance of which could influence the establishment of tetraploids in diploid populations.     

MORPHOLOGICAL AND CYTOLOGICAL VARIABILITY IN TRIPSACUM DACTYLOIDES (GRAMINEAE)

A total of 235 accessions of Tripsacum dactyloides (L.) L. representing natural populations from 88 locations distributed throughout ten states of the United States were collected in the wild and studied morphologically and cytologically. Multivariate analysis of variance (MANOVA) was applied to 17 morphological characters recorded for 868 specimens from the 88 locations, which were further combined into 25 groups to facilitate analysis. Multivariate F ratios for six comparisons involving different groups were highly significant, indicating that the populations were significantly different on the basis of all 17 characters. Non-significant MANOVA F ratios for two comparisons between sympatric diploid and polyploid accessions showed them to be morphologically similar. Discriminant function (DF) analysis was employed to determine which characters contributed towards group discrimination. No one variable was found to distinguish between groups; rather, a combination of characters was necessary, and the combination changed with each comparison. Accessions were mainly either diploid (2n = 36) or tetraploid (2n = 72), with two triploids (2n = 54) reported from separate locations in Texas. Cytological data revealed that diploids and tetraploids occupy a more sympatric distribution than previously supposed. Morphological analysis suggested that diploids could not always be distinguished from polyploids on the basis of morphology alone, unless the appropriate characters were used. Information from cytological and morphological data showed Tripsacum dactyloides to be composed of many populations covering a wide range of variability, with each population possessing a characteristic combination of morphological attributes. It was concluded that T. dactyloides can be most meaningfully analyzed in terms of particular geographic regions, rather than attempting to elucidate natural relationships by correlating data from the entire United States.     

Electrophoretic relationships and phylogeny of Nordic polyploids inDraba (Brassicaceae)

104 populations of 15 Nordic species (2x–16x) of the taxonomically complex genusDraba were investigated using enzyme electrophoresis. The polyploids were genetic alloploids showing high levels of fixed heterozygosity and electrophoretic variation; the diploids were homozygous and genetically depauperate. Thus, the data suggest that alloploidy in arctic-alpineDraba serves as an escape from genetic depauperation caused by inbreeding at the diploid level. Although some populations probably have local alloploid origins, electrophoretic data indicate that several polyploids have migrated repeatedly into the Nordic area.Draba crassifolia (2n = 40) is probably octoploid based on x = 5. A hypothesis on the evolutionary history of the polyploids based on x = 8 is presented. Diploids contributing to numerous polyploid genomes and multiple origins of polyploids have seriously blurred taxonomic relationships. Relationships inferred from genetic data do not always correspond to those based on morphology; two morphologically very similar polyploids,D. alpina andD. oxycarpa, were, for example, genetically distant and probably represent independent lineages.     

Polyploid evolution and biogeography in Chelone (Scrophulariaceae): morphological and isozyme evidence

Chelone is a genus of perennial herbs comprising three diploid species (C. cuthbertii, C. glabra, and C. lyonii) and a fourth species (C. obliqua) that occurs as tetraploid and hexaploid races. To assess patterns of isozyme and morphological variation, and to test hypotheses of hybridization and allopolyploidy, we analyzed variation among 16 isozyme loci from 61 populations and 16 morphological characters from 33 populations representing all taxa and ploidy levels. Based on morphological analyses using clustering (unweighted pair group method using an arithmetic average) and ordination (principal components analysis and canonical variance analysis) methods, we recognize three diploid species without infraspecific taxa. Polyploids in the C. obliqua complex were most similar morphologically to diploid populations of C. glabra and C. lyonii. Patterns of isozyme variation among polyploids, which included fixed heterozygosity and recombinant profiles of alleles present in diploids, suggested polytopic origins of tetraploids and hexaploids. Our data indicate independent origins of polyploids in or near the southern Blue Ridge, Interior Highlands and Plains, and Atlantic Coastal Plain regions from progenitors most similar to C. glabra and C. lyonii. Extant tetraploids were not implicated in evolution of hexaploids, and plants similar to C. cuthbertii appeared unlikely as diploid progenitors for polyploids. We propose multiple differentiation and hybridization/polyploidization cycles in different geographic regions to explain the pattern of allopatry and inferred polytopic origins among polyploids.     

Life history variation among low-arctic clones of obligately parthenogenetic Daphnia pulex: a diploid-polyploid complex

Summary Laboratory life table experiments were conducted using nine clones of obligately parthenogenetic Daphnia pulex that were collected from a site in the Canadian low-arctic. Two of the nine clones were diploids, while the other seven clones were polyploids. Significant clonal differences in age at first reproduction, size at first reproduction, number of offspring in each of the first three broods, offsrring sizes for the first two broods, and intrinsic rates of natural increase were detected. Differences in life histories were evident between polyploids and diploids. Generally, polyploid clones reached maturity at later ages, matured at larger sizes, produced smaller broods, and larger offspring than the diploid clones. The data are discussed in reference to potential biotic (i.e. invertebrate predation) and abiotic factors (i.e. physicochemical gradients) that may influence life history variation in this clonal assemblage.     

Multiple origins of polyploidy and the geographic structure of Heuchera grossulariifolia

Multiple origins of polyploidy from an ancestral diploid plant species were investigated using restriction site polymorphism and sequence variation in the chloroplast DNA (cpDNA) of Heuchera grossulariifolia (Saxifragaceae). Phylogenetic analysis indicated that autopolyploidy has arisen at least twice in the evolutionary history of this species and potentially up to as many as seven times. These results suggest a greater range of independent polyploid origins as compared to a previous study of H. grossulariifolia using cpDNA restriction sites that indicated a minimum of three independent origins. Moreover, most polyploid populations did not contain cpDNA haplotypes from a single origin, but rather combined haplotypes from at least two polyploid origins. Past migration among polyploid populations of independent origin or localized polyploid formation may explain the distribution of polyploid haplotypes within and among populations. The analysis also revealed a discrepancy between relatedness and geographical location. In nearly all sympatric populations of diploids and polyploids, polyploids had the same cpDNA haplotypes as diploids from a geographically remote population. This geographical discordance has several possible explanations, including small sample sizes, extinction of parental diploid haplotypes, chloroplast introgression, and homoplasy in the cpDNA sequence data. We conclude that the recurrent formation of polyploids is an important evolutionary mechanism in the diversification of H. grossulariifolia .     

Phylogenetic relationships and gene flow between sympatric diploid and tetraploid plants ofDactylis glomerata (Gramineae)

Phylogenetic relationships between sympatric, morphologically indistinguishable diploid and tetraploid plants ofDactylis glomerata L. (Gramineae) in Galicia (Spain) were assessed using allozyme markers for 6 distinct systems. The study exploited recent introduction in Galicia and subsequent hybridization of an alien 4xDactylis subspecies possessing distinct allozymes from those of all the native plants. Opportunities for gene exchanges between the ploidies were estimated from in situ observations of flowering, examination of progenies in 2x/4x natural and experimental crosses, and enzyme analyses. Results show a high genetic similarity between the Galician diploids and tetraploids, which possess peculiar alleles in common. Although the ploidy levels usually have distinct flowering periods, interploidal crosses do occasionally occur. Gene flow is likely much more important from the diploid to the tetraploid level. A good genetic intermixing occurs between the Galician and the alien tetraploid entities which have simultaneous flowering. Autopolyploidization of the diploids followed by various rates of hybridization is proposed as one very probable origin of natural tetraploids inDactylis.     

Habitat differentiation, hybridization and gene flow patterns in mixed populations of diploid and autotetraploid Dactylorhiza maculata s.l. (Orchidaceae)

Detailed ecological, morphological and molecular analyses were performed in mixed populations of diploid and autotetraploid Dactylorhiza maculata s.l. in Scandinavia. Comparisons were made with pure populations of either diploid ssp. fuchsii or tetraploid ssp. maculata. It was shown that mixed populations are the result of secondary contact between ssp. fuchsii and ssp. maculata. No patterns of recent and local autopolyploidization were found. Morphology and nuclear DNA markers (internal transcribed spacers of nuclear ribosomal DNA) showed that diploids and tetraploids from mixed populations have similar levels of differentiation to diploids and tetraploids from pure populations. Vegetation analyses, as well as analyses of environmental variables, revealed that diploid and tetraploid individuals in mixed populations are ecologically well differentiated on a microhabitat level. Diploids and tetraploids in pure populations have wider ecological amplitudes than they do in mixed populations. Triploid hybrids grew in intermediate microhabitats between diploids and tetraploids in the mixed populations. Plastid DNA markers indicated that both diploids and tetraploids may act as the maternal parent. Based on morphology and nuclear markers triploids are more similar to tetraploids than to diploids. There were indications of introgressive gene flow between ploidy levels. Plastid markers indicated that gene flow from diploid to tetraploid level is most common, but nuclear markers suggested that gene flow in opposite direction also may occur. Similar patterns of differentiation and gene flow appeared in localities that represented contrasting biogeographic regions. Disturbance and topography may explain why hybridization was slightly more common and the differentiation patterns somewhat less clear in the Scandinavian mountains than in the coastal lowland. An erratum to this article can be found at