The evolution of self-fertility inCrepis tectorum (Asteraceae)

The extent of self-fertility was examined in 16 populations ofCrepis tectorum. A hypothesis that a weedy habit favours autogamy was only partly supported. Low levels of self-fertility characterized non-weedy populations from calcareous grasslands (alvars) on the Baltic island in Öland. By contrast, plants in nearly all weed populations studied were more or less self-fertile. However, the trend towards autogamy may have occurred independently of the trend towards a weedy habit, as shown by moderately to high levels of self-fertility in alvar populations from two other Baltic islands. In the weed group, there was a tendency for plants from two field populations to be more autogamous than plants from more ruderal habitats. There was an association between self-fertility and small, inconspicious heads in the alvar group but the association was weaker when weed populations were also considered. The relatively wide heads characterizing the ruderal weed populations may, at least partly, be an indirect effect of increases in overall plant size and/or in the size of the fruit associated with each flower.     

Morphological responses to nutrient availability in four clonal herbs

This paper examines morphological plasticity of clonal plants of contrasting habitats and of contrasting architectures in response to nutrient supply. The hypotheses were tested that plants from rich habitats possess greater plasticity in response to variation in resource supply than species from poor habitats, and that rhizomatous species are less plastic in their response than stoloniferous species. Two sympodial rhizomatous herbs (Carex flacca, C. hirta) and two monopodial stoloniferous herbs (Trifolium fragiferum, T. repens) were subjected to four levels of nutrient supply in a garden experiment. One of the two species of each genus (C. hirta, T. repens) is from fertile and the other from infertile habitats. We measured 1) whole plant characters: total plant dry weight, number of modules (product of a single apical meristem) and number of ramets; 2) ramet characters: ramet leaf area and ramet height; and 3) spacer characters: branches per module, length per module and length per module internode.All measured characters in the Trifolium species significantly responded to treatment: the values for all measured characters increased with higher levels of fertilization. The differences in plant characters between fertilization levels were larger in Trifolium repens than in T. fragiferum in terms of whole plant characters, ramet characters and stolon internode length. The two Carex species did not differ in their responses to treatment in terms of most characters measured. In ramet characters and in some whole plant characters the species from fertile habitats were more plastic than those from infertile habitats. In spacer characters this pattern was not found. Foraging could not be demonstrated unequivocally.Morphological plasticity in the stoloniferous (Trifolium) species was much larger than in the rhizomatous (Carex) species. This seems in accordance with a foremost storage function of rhizomes, as against a foremost explorative function of stolons.     

Phenotypic plasticity of invasive Alternanthera philoxeroides in relation to different water availability, compared to its native congener

Phenotypic plasticity and genetic differentiation are two possible mechanisms that plants use to cope with varying environments. Although alligator weed (Alternanthera philoxeroides) possesses very low genetic diversity, this alien weed has successfully invaded diverse habitats with considerably varying water availability (from swamps to dry lands) in China. In contrast, its native congener (Alternanthera sessilis) has a much narrower ecological breadth, and is usually found in moist habitats. To understand the mechanisms underlying the contrasting pattern, we performed a greenhouse experiment to compare the reaction norms of alligator weed with those of its native congener, in which water availability was manipulated. Our results revealed that the two congeners had similar direction of phenotypic plasticity. However, A. philoxeroides showed greater plasticity in amount than did A. sessilis in many traits examined during the switch from wet to drought treatment. Nearly all of the phenotypic variance in A. philoxeroides could be ascribed to plasticity, while A. sessilis had a much higher fraction of phenotypic variance that could be explained by genotypic variation. These interspecific differences in plastic responses to variable water availability partially explained the difference in spatial distribution of the two congeners.     

Life-history variation in Crepis tectorum (Asteraceae)

Summary Populations of the monocarpic plant Crepis tectorum were grown in a series of uniform environments to test the hypothesis that weedy populations are more r-selected than populations from a more natural habitat. Weedy populations exhibited a combination of r- and K-selected traits. The relatively rapid growth, the potential for a summer annual habit, and the relatively high fecundity that characterized at least one of the two weed populations studied were considered as r-selected traits favored in habitats of unpredictable duration. However, high levels of competition from other weedy species or from the crop in arable fields may explain at least some presumably K-selected traits observed in the weedy populations, e.g. relatively large seeds and late flowering in the summer. Results indicated that stress due to abiotic factors (strong winds, desiccation and nutrient deficiency) has been a more important selective factor than r- or K-selection, in non-weedy populations from calcareous grasslands (alvars) on the Baltic islands.     

A phenetic study of Crepis tectorum in Fennoscandia and Estonia

Morphological variation in Crepis tectorum in Fennoscandia and Estonia was assessed in a multivariate study of herbarium material. The variation pattern revealed was more complex than that detected in earlier studies of the species in this area. There was a gradual transition between the common weed form and a similar form occurring on rocky outcrops throughout the area, particularly along the coasts and on steep hillsides on the mainland. The two endemic alvar forms on the Baltic islands of Oland and Gotland were distinct from each other but overlapped slightly with other groups. An additional outcrop form from the coast of South Finland graded into the Oland alvar form while the alvar form from Gotland was more similar to the widespread forms on the mainland of Fennoscandia and in Estonia. An outgroup comparison suggested that the widespread forms on the mainland have retained a more "ancient" morphology than the geographically restricted Baltic forms. Morphological and distributional data are used to test and extend earlier hypotheses regarding the evolution of C. tectorum in these areas. An eastern route seems likely for the first immigrations of the species into Fennoscandia. I suggest that the Baltic forms and at least some populations of the weed form were derived from non-weedy outcrop populations on the mainland, the first group when the Baltic lowland areas emerged from the sea, and the second group when man created suitable conditions for weed colonizations throughout Fennoscandia. Taxonomic implications of the results are also discussed.     

Reproductive ecology of a cleistogamous annual,Impatiens noli-tangere L., occurring under different environmental conditions

Demographic and reproductive schedules were compared among five populations of a cleistogamous annual,Impatients noli-tangere L., occurring in habitats with contrasting moisture and/or light environments. In all populations, flowering extended for 2–3 months during which light environment and mortality changed. Seasonal patterns of growth and mortality were significantly different among the five populations studied. The beginning and duration of flowering and the ratio of chasmogamous flowers to cleistogamous ones were also significantly different among populations. An experiment was conducted under different light conditions (open and closed) to separate the genetic and environmental components of the variation in reproductive traits observed among populations ofI. noli-tangere. Transplanted plants showed significant among-population variation in flowering time, as is observed in natural populations, suggesting genetic differentiation among populations of this species. On the other hand, the ratio of chasmogamous flowers did not differ among plants transplanted from three populations. Based on these results, the authors suggest that facultative cleistogamy is a conditional strategy under seasonally changing environments.     

Effects of simulated herbivory and resource availability on the invasive plant, Alternanthera philoxeroides in different habitats

In biological control programs, the insect natural enemy’s ability to suppress the plant invader may be affected by abiotic factors, such as resource availability, that can influence plant growth and reproduction. Understanding plant tolerance to herbivory under different environmental conditions will help to improve biocontrol efficacy. The invasive alligator weed (Alternanthera philoxeroides) has been successfully controlled by natural enemies in many aquatic habitats but not in terrestrial environments worldwide. This study examined the effects of different levels of simulated leaf herbivory on the growth of alligator weed at two levels of fertilization and three levels of soil moisture (aquatic, semi-aquatic, and terrestrial habitats). Increasing levels of simulated (manual) defoliation generally caused decreases in total biomass in all habitats. However, the plant appeared to respond differently to high levels of herbivory in the three habitats. Terrestrial plants showed the highest below–above ground mass ratio (R/S), indicating the plant is more tolerant to herbivory in terrestrial habitats than in aquatic habitats. The unfertilized treatment exhibited greater tolerance than the fertilized treatment in the terrestrial habitat at the first stage of this experiment (day 15), but fertilizer appears not to have influenced tolerance at the middle and last stages of the experiment. No such difference was found in semi-aquatic and aquatic habitats. These findings suggest that plant tolerance is affected by habitats and soil nutrients and this relationship could influence the biological control outcome. Plant compensatory response to herbivory under different environmental conditions should, therefore, be carefully considered when planning to use biological control in management programs against invasive plants.     

异质生境对二型花柱植物雪地报春花性状分化的影响

该研究以生长在新疆北部的二型花柱植物雪地报春(Primula nivalis)为研究材料,对7个低海拔(1 657~2 013m)森林居群和7个高海拔(2 400~2 700m)草原居群的花型比例进行调查,对花部特征进行检测,并对长、短花柱植株在不同海拔居群的花性状分化进行了比较研究,以揭示该类群植物的演化与环境的关系。结果表明:(1)在高海拔草原居群中长花柱植株比例高于短花柱植株。(2)比较草原和森林生态系统中的雪地报春居群,长花柱花形态特征间存在显著差异,但短花柱花形态特征间的差异不显著。(3)高海拔草原居群中长花柱花的花冠筒长度、雄蕊长度、雌雄蕊距离、雌雄异位程度以及雌雄蕊空间位置是导致花性状分化的主要因素。(4)在草原居群中,长花柱花的雄蕊、雌雄异位程度及花冠口至雄蕊距离的变化减少了长短花柱花间的互补式对应性。研究认为,在异质生境中长花柱花性状的变化是导致该物种长短花柱花性状分化的主要因素。     

Biocontrol of a perennial legume,Sesbania punicea,using a florivorous weevil,Trichapion lativentre: weed population dynamics with a scarcity of seeds

Summary The establishment in South Africa of a florivorous, apionid weevil, Trichapion lativentre, on Sesbania punicea, a leguminous weed of South American origin, has reduced seed production of the plants by >98%. Surveys of the age structure and density of plants in infestations of S. punicea throughout South Africa have shown that the rate of recruitment of seedlings has drastically declined within a few years in many areas, due to the weevils. However, there has unexpectedly not been a corresponding decline in the density of mature plants in extant infestations of S. punicea. In spite of this, T. lativentre has curtailed the rate of spread of the weed into uninvaded habitats and has impeded reinvasion into areas cleared of infestations by mechanical means or by another complimentary biocontrol agent.     

Experimental ecology of Dryas octopetala ecotypes

Summary Growth, survival and reproduction of adult plants from three reciprocally-transplanted populations of Dryas octopetala were followed over a seven year period, from 1979 to 1986. The two most different ecotypes, from snowbed and fellfield environments, were strongly selected against in extreme foreign environments via differential growth, survival, and reproduction. The more phenotypically-plastic ecotype from the snowbed experienced rapid, early mortality on the fellfield, prior to morphological adjustment, but no mortality in the last five years of the experiment. The less plastic fellfield ecotype showed no mortality for the first three years followed by a steady decline in numbers for the next four years. The ecotypic difference in plasticity may account for the contrasting mortality patterns. A population of plants with intermediate morphology was intermediate in fitness in extreme environments, but showed no superiority in its home site, suggesting that it is not a locally-adapted population. Natural selection maintains genetic differences between snowbed and fellfield populations in the face of gene flow. However, based on the response of intermediates, there appear to be limits to the degree to which selection can fine-tune adaptation along environmental gradients.     

Aestival and autumnal vicariads ofGentianella (Gentianaceae): A myth?

Plant samples were collected from 13 populations of the two speciesGentianella germanica andG. campestris. In addition, two samples from populations of grazed plants lacking their main stems were collected. A number of relevant morphological characters were measured, the flowering time, and the habitats of each population were studied. Some ecotypes ofG. germanica with clear differences in flowering time were recognized. Differences in flowering time and morphology between populations ofG. campestris seem to be clinal. The generally accepted theory of seasonal dimorphism with the assumption of separate aestival and autumnal vicariads proves to be inadequate. A hypothesis based upon the effect of human influences on the origin of this ecotypes is suggested.     

Population structure, genetic diversity and host specificity of the parasitic weedStriga hermonthica (Scrophulariaceae) in Sahel

Seed proteins of individual plants from 14 populations ofStriga hermonthica growing on sorghum, millet, maize and wild grasses in Burkina Faso, Mali and Niger were studied using gel electrophoresis in order to assess the population structure and genetic diversity of the parasitic weed. The relative intensity of the different bands within a pattern was used to depict the genotypic constitution of each sample. Genotypic frequencies conformed to Hardy-Weinberg expectations in 13 populations out of 14 for the two loci that were interpreted. Heterozygote deficiencies could be the result of the Wahlund effect. The genetic divergence between populations appears to be low for bothAdh andGot loci. Thus, the physiological specialization for a particular host could be a recent phenomenon. A low host specificity ofS. hermonthica populations could affect the efficiency of introducing new resistant cultivars as a control measure against the parasitic weed.     

Phenotypic plasticity rather than locally adapted ecotypes allows the invasive alligator weed to colonize a wide range of habitats

Both phenotypic plasticity and locally adapted ecotypes may contribute to the success of invasive species in a wide range of habitats. Here, we conducted common garden experiments and molecular marker analysis to test the two alternative hypotheses in invasive alligator weed (Alternanthera philoxeroides), which colonizes both aquatic and terrestrial habitats. Ninety individuals from three pairs of aquatic versus terrestrial populations across southern China were analyzed, using inter simple sequence repeat (ISSR) marker, to examine population differentiation in neutral loci. Two common gardens simulating aquatic and terrestrial habitats were set up to examine population differentiation in quantitative traits. We found no evidence of population differentiation in both neutral loci and quantitative traits. Most individuals shared the same ISSR genotype. Meanwhile, plants from different habitats showed similar reaction norms across the two common gardens. In particular, plants allocated much more biomass to the belowground roots in the terrestrial environment, where alligator weed may lose part or all of the aboveground shoots because of periodical or accidental disturbances, than those in the aquatic environment. The combined evidence from molecular marker analysis and common garden experiments support the plasticity hypothesis rather than the ecotype hypothesis in explaining the adaptation of alligator weed in a wide range of habitats.     

Morphometric differentiation,patterns of interfertility,and the genetic basis of character evolution inCrepis tectorum (Asteraceae)

Morphometric analyses of populations ofCrepis tectorum raised under uniform conditions support the earlier finding based on herbarium studies that extensive divergence has taken place in western Europe with a centre of diversity in the Baltic lowland area. A crossing experiment showed that subsp.pumila on the island of Öland has acquired a wide range of derived characters without the establishment of strong reproductive barriers, the only exception being a weak tendency for crosses with other populations on Öland to yield more fertile F₁ offspring than crosses with weedy and non-weedy populations in adjacent regions. Canadian weed populations showed a more uniform response in crosses with subsp.pumila than did morphologically similar weed populations scattered throughout Central Europe. While F₂ data suggest a monogenic basis of differences in the extent of leaf dissection, other distinctive traits appear to be governed by multiple genetic factors with individually slight effects.     

Morphometric analysis of the genusHemerocallis L. (Liliaceae) in Korea

To better understand the patterns of variability and distributions ofHemerocallis in Korea, 53 locations were visited and measurements of 19 morphological and phenological characters were taken on plants directly from their natural habitats. For morphometric analysis, 10 plants from each of 34 populations and five herbarium specimens ofH. middendorffii were used and the data from 12 quantitative characters was analyzed using univariate analysis. Except the littoral populations of Cheju, Hong, Taehuksan, and Sohuksan Islands (H. hongdoensis M. Chung & S. Kang), three peninsular KoreanHemerocallis species can be recognized mainly in South Korea:H. hakuunensis Nakai (=H. micrantha Nakai, growing on southern, central, and northwestern Korea);H. thunbergii Baker (=H. coreana Nakai, found on southeastern and central Korea); andH. middendorffii Tr. et Mey. (central and northeastern Korea). Morphological and phenological features contributing to recognition of the three groups were; color of perianth, shape of roots, shape of inflorescence, flowering time, odor, length of inflorescence, width of the lowest bracts, length of perianth tube enclosing a ovary, width of the inner perianth lobes. Natural hybridization seems to be rare in KoreanHemerocallis. It appears that the KoreanHemerocallis species are relatively well characterized by their distribution patterns, phenology, and habitats compared with the JapaneseHemerocallis species.     

Establishment and dispersal of the biological control weevil Rhinoncomimus latipes on mile-a-minute weed, Persicaria perfoliata

Mile-a-minute weed, Persicaria perfoliata (L.) H. Gross (Polygonaceae), is an annual vine from Asia that has invaded the eastern US where it can form dense monocultures and outcompete other vegetation in a variety of habitats. The host-specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the US in 2004 as part of a classical biological control program. The weevil was intensively monitored in three release arrays over 4 years, and field cages at each site were used to determine the number of generations produced. The weevil established at all three sites and produced three to four generations before entering a reproductive diapause in late summer. Weevils dispersed at an average rate of 1.5–2.9 m wk⁻¹ through the 50 m diameter arrays, which had fairly contiguous mile-a-minute cover. Weevils dispersing in the broader, more variable landscape located both large monocultures and small isolated patches of mile-a-minute 600–760 m from the release within 14 months. Weevil density ranged from fewer than 10 to nearly 200 weevils m⁻² mile-a-minute weed. Mile-a-minute cover decreased at the site with the highest weevil density. The production of P. perfoliata seed clusters decreased with increasing weevil populations at two sites, and seedling production declined over time at two sites by 75% and 87%. The ability of the weevil to establish, produce multiple generations per season, disperse to new patches, and likelihood of having an impact on plants in the field suggests that R. latipes has the potential to be a successful biological control agent.     

Buffering effect of B-chromosome system of Trigonella foenum-graecum against different soil types

Summary Accessory (B–) chromosomes have been reported in many plants in various regions of the world. Studies on natural populations of certain plants like Phleum phleoides revealed that B-chromosomes were present in different frequencies in different habitats indicating a correlation between B-frequency and environmental conditions. The present experiment was performed to determine the response of B-chromosome carrier and non-carrier plants of Trigonella to different soil conditions. When the morphological data was subjected to analysis of variance, it was found that the carrier plants exhibited uniformity of characters in the different soil types, while the non-carrier plants exhibited considerable variability. The conclusion that can be drawn from this experiment is that B-chromosomes act as a buffering agent, neutralising the variability effect of different soil types.     

Origin matters for level of resource sharing in the clonal herb <Emphasis Type="Italic">Aegopodium podagraria</Emphasis>

Resource sharing in heterogeneous environments has been shown to increase growth and survival of clonal plants. In addition, plants in harsh climates have been suggested to have higher levels of resource sharing than plants in milder climates. We experimentally investigated the level of resource sharing in plants from garden and forest habitats from two regions with contrasting climates. The clonal herb Aegopodium podagraria reaches its northern distribution limit in central Sweden. South of that it grows in both patchy and dynamic light climate, as a natural component of deciduous forest, as well as in more homogeneous light climates as a garden weed. Since heterogeneity and habitat harshness have both been suggested to increase resource sharing, we hypothesized (1) integration and sharing of resources to be higher in plants from forest than in plants from garden habitats and (2) integration and sharing of resources to be higher in plants from the northern region that encounter a harsher climate than in plants from the southern region. Clonal fragments of A. podagraria were collected and multiplied in the greenhouse. Ramet pairs were then planted in adjacent pots, with one ramet shaded. Rhizome connection was either left intact or severed to prevent resources sharing. Plants from forest habitats were more negatively affected by the severance treatment than plants from garden habitats. Although region alone had no significant effect on biomass, the interaction between rhizome severing, shading and regional origin was close to significance. We conclude that A. podagraria from forests are more dependent on resource sharing than those from gardens. These results concur with previous studies that suggest that local adaptation for different degrees of resource sharing can occur in clonal plants.     

Geographical variation and genetic analysis of leaf shape inCrepis tectorum (Asteraceae)

Canonical variate analysis of plants raised in a uniform environment was used to study the pattern of geographical variation in leaf shape ofCrepis tectorum (Asteraceae). The diversity in leaf shape was much greater among populations confined to areas with exposed bedrock in the Baltic region than among weed populations scattered throughout Europe and Canada. A Ward's clustering linked outcrop populations from the archipelago of SW. Finland and the islands of Öland (Sweden) and Saaremaa (Estonia) due to the deeply lobed leaves characterizing these populations, while outcrop populations along the coast of E. Sweden were grouped due to their weakly lobed, narrow and dentate leaves. Most of the weed populations were grouped together but there was no tendency for the variation in this group to be related to habitat or geographical location. A mosaic of variation reflected in sharp (random) differentiation among local populations was superimposed on the large-scale ecogeographical pattern.—Crossing data indicated that most of the variation in degree of leaf dissection is governed by one major gene with deeply lobed leaves dominant over weakly lobed leaves. I suggest that the simple pattern of inheritance may have favoured rapid evolutionary changes in leaf shape, particularly in the Baltic area which emerged relatively late from the sea. Genetic correlations may have constrained the pattern of variation at higher taxonomic levels, since some of the trait associations detected in a segregating F₂ generation were also found at the among-population level.     

Life history variation and infraspecific heterochrony inRhinanthus glacialis (Scrophulariaceae)

Comparative cultivation experiments were made with 14 populations of the annual, hemiparasitic speciesRhinanthus glacialis from seven different habitat types. Phenological data were recorded and plants from field samples and from corresponding experimental samples were morphologically analysed.There are up to ten weeks difference in the start of flowering between means of the extreme experimental populations. Flowering time variation correlates with the number of internodes and also with most of the other morphological characters. Flowering time also shows close relationships to different grassland types and the examined populations appear to be well-adapted to the environmental conditions in natural or man-influenced grassland types as far as flowering time is concerned. The number of internodes is mostly insensitive to the applied experimental influences. The rate of internode production is constant. Trade-offs are suggested for some morphological traits (i.e., few internodes with few branches and thus few flowers as trade-off for early flowering). — Previous studies of life history variation based on population data have not been incorporated into the theory of heterochrony. The pattern of life history variation of populations ofR. glacialis together with information about morphology, ecology, and phylogeny suggests that some variants have evolved from ancestors via heterochronic processes. Earlier flowering populations from montane hay meadows, litter meadows, and dry meadows belong to paedomorphic variants. These have fewer internodes and branches as well as reduced height in comparison with the suggested ancestor. Later flowering populations from subalpine limestone grassland belong to a peramorphic variant with more internodes, more branches, and taller stems in comparison with the suggested ancestor.