Early evolution in a hybrid swarm between outcrossing and selfing lineages in Geum

Although often considered as evolutionary dead ends, selfing taxa may make an important contribution to plant evolution through hybridization with related outcrossing lineages. However, there is a shortage of studies examining the evolutionary dynamics of hybridization between outcrossing and selfing taxa. On the basis of differential pollinator attractiveness, production and competitive ability of pollen, as well as levels of inbreeding depression, we predict that the early products of hybridization between outcrossing and selfing lineages will be F1s and first-generation backcrosses sired mainly by the outcrossing lineage, together with selfed F2s containing a limited genetic contribution from the outcrosser. These predictions were tested using amplified fragment length polymorphism and chloroplast markers to analyze the composition of a recent hybrid swarm between predominantly outcrossing Geum rivale and predominantly selfing Geum urbanum. In line with predictions, the hybrid swarm comprised both parental species together with F1s and first-generation backcrosses to G. rivale alone. Chloroplast data suggested that G. rivale was the pollen parent for both observed hybrid classes. However, there was no evidence for F2 individuals, despite the fact that the F1 was fully self-compatible and able to auto-pollinate. The pollen fertility of F1s was only 30% lower than that of the parental taxa, and was fully restored in backcross hybrids. Predicting future evolution in the hybrid swarm will require an understanding of the mating patterns within and among the mix of parental, F1 and backcross genotypes that are currently present. However, these results support the hypothesis that introgression is likely to be asymmetrical from selfing to outcrossing lineages.     

Interaction between perch (Perca fltuviatilis), and brown trout, (Salmo trutta)

A decline in the growth rate of perch in Malham Tarn, Yorkshire, has occurred in recent times. This was found to coincide precisely with the resumption of stocking of brown trout, after a period of several years during which no stocking took place. In the absence of any other known correlating factor, it is suggested that the increase in population density of the trout may be responsible for the observed decline in perch growth.     

Landscape genetics of the self-compatible forest herb Geum urbanum: effects of habitat age, fragmentation and local environment

To investigate the role of habitat fragmentation, fragment age and local environment in shaping the genetics of plant populations, we examined the genetic structure of the self-compatible forest herb Geum urbanum using microsatellite markers. A historical land-use reconstruction assigned the studied populations to two age classes: populations in primary forest fragments, and populations in secondary fragments. Local environmental conditions were quantified on the basis of the herb-layer community composition. A stepwise general linear model revealed that levels of within-population genetic diversity were best explained by population size, landscape connectivity and the interaction between both. Connectivity was positively correlated with the genetic diversity of small populations, but did not significantly affect the diversity of large populations. Contrary to what we expected, secondary-forest populations showed lower divergence relative to populations located in primary patches. Small populations were genetically more diverged compared to large populations. Mantel tests showed no significant isolation by distance and no significant correlation between habitat similarity and genetic differentiation. We conclude that gene flow has probably prevented founder events from being reflected in the present genetic structure of G. urbanum. Gene flow towards low-connectivity populations, however, seemed to be insufficient to counteract the effects of drift in small populations.     

Factors affecting the distribution of some Phryganeaeid (Trichoptera) in Malham Tarn, Yorkshire

This paper describes a distribution problem involving Agrypnetes crassicornis and Phryganea obsoleta (Trichoptera : Phryganeidae) in Malham Tarn, Yorkshire. The only suitable habitat for their larvae is the offshore vegetation, chiefly Chara delicatula, but owing to the specialized egglaying behaviour of this family and the absence of other sites, the eggs of Agrypnetes crassicornis are laid on the exposed rocky shoreline. The situation with Phryganea obsoleta differs in that the emergence of Potamogeton spp. provides new oviposition sites away from the shore. Investigations on both species however have provided further support for an hypothesis which postulates the importance of wind-induced water movements in rapidly dispersing newly hatched insect larvae to the favourable habitats in deep water.     

Taxonomic notes on Geum micropetalum Gasp. and Geum heterocarpum Boiss. (Rosaceae) in Italy

The systematic and taxonomic position of Geum micropetalum, a controversial species currently treated as endemic to Italy, is discussed. The name G. micropetalum is here typified and shown to be a heterotypic synonym of G. urbanum. Herbarium survey evidenced the historical presence in Italy of G. heterocarpum. During ad hoc field investigations carried out in 2012, we were able to confirm the presence of this species in a single small population, the only one known from Italy.     

Allelopathic inhibition of germination by Alliaria petiolata (Brassicaceae)

Garlic mustard (Alliaria petiolata, Brassicaceae) is an invasive, nonindigenous species currently invading the understory of North American woodlands where it is a serious threat to the native flora. Part of this success might be due to allelopathic interference by garlic mustard. Two congeneric species, the European Geum urbanum and the North American Geum laciniatum, were tested for allelopathic inhibition of germination by garlic mustard. Seeds were germinated either on substrate contaminated by garlic mustard or on substrate with contamination neutralized by activated carbon. Allelopathic effects of native European and invasive North American garlic mustard populations were also compared. Activated carbon increased germination by 14%, indicating that garlic mustard contaminated the substrate through root exudates. Activated carbon in turn counteracted this effect. The two test species differed in their sensitivity to allelopathic interference. North American G. laciniatum had a much stronger increase in germination when activated carbon was added to the substrate, independent of the origin of garlic mustard. In contrast, the European G. urbanum germinated better in substrate precultivated with North American garlic mustard, whereas activated carbon increased its germination only in substrate precultivated with European garlic mustard.     

Maintaining their genetic distance: Little evidence for introgression between widely hybridizing species of Geum with contrasting mating systems

Within the plant kingdom, many genera contain sister lineages with contrasting outcrossing and inbreeding mating systems that are known to hybridize. The evolutionary fate of these sister lineages is likely to be influenced by the extent to which they exchange genes. We measured gene flow between outcrossing Geum rivale and selfing Geum urbanum, sister species that hybridize in contemporary populations. We generated and used a draft genome of G. urbanum to develop dd‐RAD data scorable in both species. Coalescent analysis of RAD data from allopatric populations indicated that the species diverged 2–3 Mya, and that historical gene flow between them was extremely low (1 migrant every 25 generations). Comparison of genetic divergence between species in sympatry and allopatry, together with an analysis of allele frequencies in potential parental and hybrid populations, provided no evidence of contemporary introgression in sympatric populations. Cluster‐ and species‐specific marker analyses revealed that, apart from four early‐generation hybrids, individuals in sympatric populations fell into two genetically distinct groups that corresponded exactly to their morphological species classification with maximum individual admixture estimates of only 1–3%. However, we did observe joint segregation of four putatively introgressed SNPs across two scaffolds in the G. urbanum population that was associated with significant morphological variation, interpreted as tentative evidence for rare, recent interspecific gene flow. Overall, our results indicate that despite the presence of hybrids in contemporary populations, genetic exchange between G. rivale and G. urbanum has been extremely limited throughout their evolutionary history.     

Population genetic diversity of the clonal plant Geum reptans (Rosaceae) in the Swiss Alps

In the alpine landscape most plant populations are spatially isolated due to extreme patchiness and strong natural fragmentation. We used RAPD-PCR (randomly amplified polymorphic DNA polymerase chain reaction) for a study of the genetic diversity within and among 20 populations of Geum reptans, an outcrossing clonal plant species in the Swiss Alps. Populations were sampled at different altitudes, in early-, medium- and late-successional habitats (population origin) using a spatially hierarchical design, with distances among populations ranging from 0.2 to 208 km. Seed and pollen dispersibility was estimated by direct measurements. Seed dispersibility by wind was low with only 0.015% of the seeds flying over 100 m. Observed pollen flow was even more restricted. Molecular diversity within populations was irrespective of population origin (H(e) = 0.22 ± 0.004) and similar to the average of other RAPD studies. Contrary to our expectation, populations were only moderately differentiated (G(st) = 0.14). However, there was a clear spatial genetic structure and a positive relationship between pairwise genetic and geographic distances. Our results indicate considerable gene flow among populations within the same regional area, and we found no indication for genetic depletion during succession or in peripheral habitats. We conclude that, despite the high natural fragmentation and the importance of vegetative reproduction in this alpine plant, gene flow and repeated seedling recruitment during succession might be more frequent than commonly suggested.     

The occurrence of the trematode Plagiorchis muris in the wood mouse Apodemus sylvaticus in North Yorkshire, UK

Wood mice Apodemus sylvaticus were trapped each September over a 13-year period, from 1993 to 2005, in a wooded area adjacent to Malham Tarn, Yorkshire, UK. Plagiorchis muris was found to be the dominant intestinal trematode and occurred in every year of sampling, with an overall prevalence of 16.9%. This appears to be the first record of P. muris in A. sylvaticus within the UK. The mean worm burden was 2.03 and the distribution of the parasite within mice was typically overdispersed. No difference in prevalence relative to host sex was evident although there was a higher prevalence of 21.9% in larger older mice compared with 9.1% in juveniles, which probably relates to a greater foraging activity in older mice. Annual prevalence values of P. muris infections varied significantly over the study period with higher prevalences being associated with years with a high spring/summer rainfall. The second intermediate hosts for P. muris include a variety of aquatic insect larvae and it is likely that the higher rainfall may result in the occurrence of temporary water bodies suitable for the development of insect larvae.     

Gastropod associations within Malham Tarn, Yorkshire

A survey of gastropod species associations was undertaken in one locus, i.e. Malham Tarn, Yorkshire. This ranged through both the major weed beds and the rocky shores. A semiquantitative analysis, in terms of both presence-absence and relative abundance indices, suggested a clear distinction between the snail fauna present in the weed beds and that on the rocky shores, and also indicated a separation of faunas within these two major habitat types. The latter was closely correlated with degree of exposure. From this, and other evidence, it was suggested that wind, via wave action, may be a critical determinant of both the qualitative and quantitative aspects of biocoenotic structure in large bodies of fresh water.     

Habitat distributional change of the large copper,Lycaena dispar (Lepidoptera: Lycaenidae), on the Korean Peninsula: Studies along the Geum River

Habitat changes of butterflies are caused by various factors, including food plants, natural enemies, and climate change. Recently, new habitats of the large copper have been reported on the Korean Peninsula, mostly along a riverside. This implies that this butterfly may move along a drainage system. In this study, new habitats of the large copper along the Geum River were investigated and the population dynamics of the butterfly were monitored in selected areas through the mark–release–recapture method. The results showed that the large copper had dispersed inland along the Geum River, which runs along the central western seashore of South Korea. In addition, the habitat distribution of the butterfly showed good coincidence with the distribution of a sorrel, Rumex japonicas, the main food plant of the large copper. A study on the moving distance of the butterfly showed that females moved further than males, while males showed more frequent movement within a short distance. The movement of the large copper along the Geum River, which lies over 100 km south of the Han River (a habitat previously reported in 1973), suggests that the movement of the butterfly might be more affected by human activities, such as the development of bike paths along the Geum River and maintenance of the riverbed, than climate change.     

Genotype, soil type, and locale effects on reciprocal transplant vigor, endophyte growth, and microbial functional diversity of a narrow sagebrush hybrid zone in Salt Creek Canyon, Utah

When addressing the nature of ecological adaptation and environmental factors limiting population ranges and contributing to speciation, it is important to consider not only the plant's genotype and its response to the environment, but also any close interactions that it has with other organisms, specifically, symbiotic microorganisms. To investigate this, soils and seedlings were reciprocally transplanted into common gardens of the big sagebrush hybrid zone in Salt Creek Canyon, Utah, to determine location and edaphic effects on the fitness of parental and hybrid plants. Endophytic symbionts and functional microbial diversity of indigenous and transplanted soils and sagebrush plants were also examined. Strong selection occurred against the parental genotypes in the middle hybrid zone garden in middle hybrid zone soil; F(1) hybrids had the highest fitness under these conditions. Neither of the parental genotypes had superior fitness in their indigenous soils and habitats; rather F(1) hybrids with the nonindigenous maternal parent were superiorly fit. Significant garden-by-soil type interactions indicate adaptation of both plant and soil microorganisms to their indigenous soils and habitats, most notably in the middle hybrid zone garden in middle hybrid zone soil. Contrasting performances of F(1) hybrids suggest asymmetrical gene flow with mountain, rather than basin, big sagebrush acting as the maternal parent. We showed that the microbial community impacted the performance of parental and hybrid plants in different soils, likely limiting the ranges of the different genotypes.     

Evaluation of the Degraded Riparian Ecosystems in the Geum River Watershed in Korea

Riparian vegetation, an important mediator of land–water interactions, provides habitat for animals and other organisms; however, riparian vegetation zones have been altered by agricultural and urban development in Korea. This riparian vegetation survey was conducted to obtain information vital for the ecological restoration and management of the Korean Geum River ecosystem. At 100 study sites, along the Geum River, we recorded the vegetation of the Geum riparian zone. We then surveyed the riparian vegetation associations in the area and overlaid those areas corresponding to trees, shrubs, perennial herbs, annual herbs, exotic plants, cultivated lands, and damaged lands on a geographical map. We also reconstructed the cross-sectional landscape. The mean values of vegetation diversity, exotic plant area (%), annual plant area (%), and species richness were 6.47 ± 0.26, 5.44 ± 1.01, 11.98 ± 1.20, and 22.69 ± 0.93, respectively. The landscape elements of the herbaceous plants were more spread out, compared with those of the woody plants, and 23 sites were composed strictly of herbs. Our results indicate significant differences in vegetation structure among the study sites. For example, at some sites, exotic plants, cultivated lands, and damaged lands dominated the landscape comprising 25.7, 62, and 68.9%, respectively, of the area. The riparian landscape reference model suggested by these results may be applied to studies of other well-conserved riparian zones. We propose that the material pathways and transport of organisms from land to water at Geum River depend on the patchy distribution of these diverse landscape elements.     

Relationships of Plant Parasitic Nematodes to Sites in Native Iowa Prairies

Soil samples were collected from three native Iowa prairies and analyzed for plant paiasitic nematodes and selected soil properties. Sites or nematodes were clustered with similarities related to habitat by a cluster analysis of site by nematode species and of nematodes by site. Some nematodes occurred in a wide range of prairie habitats, whereas others were more restricted. For example, greater numbers of Xiphinema americanum were in the low, well-drained sites than in the low wet sites or upland dry sites. Wet sites contained fewer nematodes than well-drained sites. Well-drained sites contained mainly Tylenchorhynchus maximus, Helicotylenchus pseudorobustus, and X. americanum. Wetter sites contained almost exclusively X. chambersi, H. hydrophilus, Telylenchus joctus, and an undescribed species of Tylenchorhynchus.     

Helminth parasites of the wild rabbit Oryctolagus cuniculus near Malham Tarn, Yorkshire, UK

Between 1992 and 1996, 95 rabbits from the immediate locality of Malham Tarn, North Yorkshire, UK were examined for the presence of helminth parasites. All the examinations took place in late September or October. Three species of nematodes, Graphidium strigosum, Passalurus ambiguus and Trichostrongylus retortaeformis and two species of cestodes, Taenia pisiformis and Cittotaenia pectinata were identified. There were no associations between helminth species richness and year of sampling, host weight or sex. A logistic model was fitted to the prevalence data from these helminths as was an over-dispersed Poisson model to the worm burden data. Graphidium strigosum was the most frequently identified species with an average prevalence of 78%. The mean prevalence and intensity of Graphidium infection were significantly effected by sampling year. The lower than normal rainfall recorded at the Tarn during the years 1995 and 1996 may have be one reason for this pattern. The worm burden of G. strigosum was significantly positively associated with rabbit body weight. The intensity of infection with P. ambiguus was significantly higher in female rabbits. There was a significant non-linear relationship between P. ambiguus worm burden and rabbit weight (P = 0.002) with worm burdens being highest in the 1000 g to 1499 g weight cohort. Trichostrongylus retortaeformis was only identified in 1994 and male rabbits harboured significantly higher worm burdens than females (48 vs. 7, P = 0.022). Over the five years, the average Taenia pisiformis prevalence was 31% and there was a significant positive association between worm burden and rabbit weight (P = 0.001). Cittotaenia pectinata had a prevalence of 37% over the whole study period with no interactions between prevalence or intensity and body weight, year of sampling or rabbit sex. All five helminths showed an overdispersed distribution with k values less than 1.     

Iridoid patterns in Galium L. and some phylogenetic considerations

From 19 species of Galium, members of 6 European sections of the genus, 24 compounds were isolated, namely 16 iridoid glucosides, 2 secoiridoid glucosides and 6 triterpene saponins (the later found only in G. rivale (Sibth. & Sm. Griseb.) The iridoid content was analyzed by thin layer chromatography - densitometry. An effort was made to clarify interspecies relationships, based on the obtained results and previous data. Generally, a nearly uniform iridoid pattern in the studied species was observed. Nevertheless, some distinctions gave reason the following chemical characters to be treated as taxonomic markers: iridoids, secogalioside (characteristic of G. mollugo group), iridoids V1 and V2 (G. humifusum Bieb. and G. verum L.), 6-acetylscandoside (G. incurvum group and G. verum) and the triterpene saponins, rivalioside A and rivalioside C (characteristic of G. rivale). The studied species regarding to the iridoids could be attributed to three lines of evolutionary differentiation. One line is leading to the differentiation of G. rivale. It contains specific triterpenoids as well as iridoid acids, which show parallel development of both glyceraldehyde 3-phosphate/pyruvate and mevalonate biosynthetic routes in this species. A second line includes G. mollugo and G. incurvum species groups and the species G. humifusum and G. verum. Variety of iridoid esters, hydroxy and carboxy derivatives of iridoids and secoiridoids characterised this line. Third line comprises the remaining studied species, members of different sections and species groups. They posses a nearly identical iridoid pattern, which suggests a convergent evolution regarding to the iridoids.     

Photosynthetic Adaptation to Light Intensity in Plants Native to Shaded and Exposed Habitats

Photosynthetic adaptation to light intensity has been studied in clones of populations from shaded and exposed habitats of Rumex acetosa and Geum rivale. Clones of the shade species Lamium galeobdolon and the sun species Plantago lanceolata were also included for comparison. The plants were grown under controlled conditions at a high and a low light intensity. The capacity of photosynthetic carbon dioxide uptake at low as well as at saturating light intensities was determined on single attached leaves. As was previously demonstrated in Solidago virgaurea, clones of populations native to shaded and to exposed environments show differences in the photosynthetic response to light intensity during growth. The data provide evidence that populations of the same species native to habitats with contrasting light intensities differ in their photosynthetic properties in an adaptive manner Ln a similar mode as sun and shade species.     

Analysis of somatic hybrids between two sterile dihaploid Solanum tuberosum L. breeding lines. Restoration of fertility and complementation of G. pallida Pa2 and Pa3 resistance

Fourteen somatic hybrids generated by electrofusion of mesophyll protoplasts from a non-flowering dihaploid S. tuberosum clone, DHAK-11, and a male-sterile dihaploid clone S. tuberosum, DHAK-33, were grown in the greenhouse and subjected to morphological assessments and tests for fertility and resistance to the white potato cyst nematode Globodera pallida pathotypes Pa2 and Pa3. The ploidy level of the hybrids ranged from 38 to 63 chromosomes. All hybrids developed flowers with violet petals except for one, hy-56, that possessed red petals. The colour of the tuber skin was purple in all hybrids except in hy-56 where the tuber skin was red. All of the hybrids were female fertile and generated viable seeds. Near-tetraploid hybrids produced the highest number of seeds per fruit and these seeds had a normal size. Hybrids with 58 or more chromosomes produced smaller seeds and less seeds per fruit. The germination frequency of the seeds was not influenced by the chromosome number of the hybrids. Pollen viability was determined and the male fertility of three hybrids was tested. Pollination with these three hybrids gave rise to fruit development, but only one produced viable seeds. The hybrids were tested for resistance to G. pallida pathotypes Pa2 and Pa3. A high level of resistance to Pa3, inherited from one parental clone, DHAK-11, and a high level of resistance to Pa2, inherited from the other parental clone, DHAK-33, was combined in four hybrids. These results demonstrate, that protoplast fusion is an efficient method for restoring the fertility of somatic hybrids generated from sterile parent clones, and is a powerful procedure for the complementation of multigenetic disease resistance traits in potato breeding lines.     

Population dynamics of Agrimonia eupatoria and Geum rivale,two perennial grassland species

The population dynamics of two polycarpic perennials,Agrimonia eupatoria and Geum rivale, characteristic ofsemi-natural grasslands in Scandinavia, were examined in south-eastern Sweden.The perennial forbs were studied for several years in two populations each,located in habitats characteristic for the species in the study area.Demographic transition probabilities varied significantly between populationsand among years for the species. Transition matrix modelling yielded-values (population growth rates) that ranged from 0.89 to 1.14 forA. eupatoria and from 0.94 to 1.04 for G.rivale. The elasticity analysis showed that stasis in the adultstageclasses contributed most to . The life-table response experimentanalysis produced similar results as the elasticity analysis, where stasistogether with progression in the adult stage classes made a large contributionto the observed spatial variation in . Simulations of expected time toextinction were in the order of centuries for the study populations. Seedlingrecruitment was enhanced by seed addition and small scale disturbance inpopulations. For the intermediately abundant and more patchily distributedA. eupatoria, a regional survey of local populations wasconducted in the study area which revealed that most populations wererelativelysmall (< 100 individuals) and restricted to grassland fragments in roadverges. Furthermore, an attempt was made to estimate fruit dispersal from localpopulations.     

GERMINATION ECOLOGY OF DANTHONIA SERICEA POPULATIONS

Seed were collected from nine populations of Danthonia sericea on either wet or well-drained, sandy sites ranging from Georgia to New Jersey. Germination tests were conducted in the laboratory and in field plots established on contrasting habitat types. Factors considered were temperature, light, prechilling, scarification, and after-ripening. The majority of freshly harvested seed from all nine populations exhibited innate dormancy the intensity of which decreased with age. Dormancy was removed by scarification and prechilling treatments of 60 days. Seed from wet habitats showed significantly greater germination in light (alternating night-day) than in dark, while seed from well-drained habitats had a higher germination percentage in dark than in light. Both responses are considered to be physiological adaptations of the populations to their respective habitats. Alternating night-day temperatures of 15–25 C produced the greatest overall germination for the nine populations; 15–20 and 10–25 C produced the next highest responses. Field observations confirmed that peak germination occurred in the spring, at the time soil temperatures were in the same range found to be optimum in the laboratory. A comparison of the germination responses obtained from six New Jersey populations in field plots on contrasting habitat types further revealed distinctive physiological adaptations for populations from differing habitats.