Inbreeding in Mimulus guttatus Reduces Visitation by Bumble Bee Pollinators

Inbreeding in plants typically reduces individual fitness but may also alter ecological interactions. This study examined the effect of inbreeding in the mixed-mating annual Mimulus guttatus on visitation by pollinators (Bombus impatiens) in greenhouse experiments. Previous studies of M. guttatus have shown that inbreeding reduced corolla size, flower number, and pollen quantity and quality. Using controlled crosses, we produced inbred and outbred families from three different M. guttatus populations. We recorded the plant genotypes that bees visited and the number of flowers probed per visit. In our first experiment, bees were 31% more likely to visit outbred plants than those selfed for one generation and 43% more likely to visit outbred plants than those selfed for two generations. Inbreeding had only a small effect on the number of flowers probed once bees arrived at a genotype. These differences were explained partially by differences in mean floral display and mean flower size, but even when these variables were controlled statistically, the effect of inbreeding remained large and significant. In a second experiment we quantified pollen viability from inbred and self plants. Bees were 37–54% more likely to visit outbred plants, depending on the population, even when controlling for floral display size. Pollen viability proved to be as important as floral display in predicting pollinator visitation in one population, but the overall explanatory power of a multiple regression model was weak. Our data suggested that bees use cues in addition to display size, flower size, and pollen reward quality in their discrimination of inbred plants. Discrimination against inbred plants could have effects on plant fitness and thereby reinforce selection for outcrossing. Inbreeding in plant populations could also reduce resource quality for pollinators, potentially resulting in negative effects on pollinator populations.     

THE EFFECTS OF FIVE GENERATIONS OF ENFORCED SELFING ON POTENTIAL MALE AND FEMALE FUNCTION IN MIMULUS GUTTATUS

In prior work we detected no significant inbreeding depression for pollen and ovule production in the highly selfing Mimulus micranthus, but both characters showed high inbreeding depression in the mixed-mating M. guttatus. The goal of this study was to determine if the genetic load for these traits in M. guttatus could be purged in a program of enforced selfing. These characters should have been under much stronger selection in our artificial breeding program than previously reported characters such as biomass and total flower production because, for example, plants unable to produce viable pollen could not contribute to future generations. Purging of genetic load was investigated at the level of both the population and the individual maternal line within two populations of M. guttatus. Mean ovule number, pollen number, and pollen viability declined significantly as plants became more inbred. The mean performance of outcross progeny generated from crosses between pairs of maternal inbred lines always exceeded that of self progeny and was fairly constant for each trait through all five generations. The consistent performance of outcross progeny and the universally negative relationships between performance and degree of inbreeding are interpreted as evidence for the weakness of selection relative to the quick fixation of deleterious alleles due to drift during the inbreeding process. The selective removal (purging) of deleterious alleles from our population would have been revealed by an increase in performance of outcross progeny or an attenuation of the effects of increasing homozygosity. The relationships between the mean of each of these traits and the expected inbreeding coefficient were linear, but one population displayed a significant negative curvilinear relationship between the log of male fertility (a function of pollen number and viability) and the inbreeding coefficient. The generally linear form of the responses to inbreeding were taken as evidence consistent with an additive model of gene action, but the negative curvilinear relationship between male fertility and the inbreeding coefficient suggested reinforcing epistasis. Within both populations there was significant genetic variation among maternal lineages for the response to inbreeding in all traits. Although all inbred lineages declined at least somewhat in performance, several maternal lines maintained levels of performance just below outcross means even after four or five generations of selfing. We suggest that selection among maternal lines will have a greater effect than selecting within lines in lowering the genetic load of populations.     

Genetic variation and covariation for resistance and tolerance to Cucumber mosaic virus in Mimulus guttatus (Phrymaceae): a test for costs and constraints

Genetic variation for resistance and tolerance to pathogens may be maintained by costs represented as genetic tradeoffs between these traits and fitness. The evolution of resistance and tolerance also may be constrained by negative genetic correlations between these defense systems. Using a complete diallel, we measured genetic variation and covariation for and among performance, resistance, and tolerance traits in Mimulus guttatus challenged with a generalist pathogen, Cucumber mosaic virus (CMV). Viral coat protein was detected by enzyme-linked immunosorbent assay (ELISA) in all inoculated plants, indicating that all plants were susceptible to infection, although the ELISA absorbance varied quantitatively across plants. Plants inoculated with CMV had significantly reduced aboveground biomass and flower production relative to controls, although date of first flower was unaffected by infection. All three of these performance traits showed moderate to high narrow-sense heritability (h2 = 0.32-0.62) in both inoculated and control plants. We found phenotypic variation for both tolerance of and resistance to our strain of CMV, but both displayed very low narrow-sense heritability (h2 < 0.03). We found no evidence of a trade-off between resistance and tolerance. We also found no evidence for a cost of resistance or tolerance. In fact, a significant genetic correlation suggested that plants that were large when healthy had the greatest tolerance when infected. Significant, positive genetic correlations found between performance of uninfected and infected plants suggested that selection would likely favor the same M. guttatus genotypes whether CMV is present or not.     

Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae)

Previous studies have demonstrated genetic variation for resistance to insect herbivores and host plant quality. The effect of plant mating system, an important determinant of the distribution of genetic variation, on host plant characteristics has received almost no attention. This study used a controlled greenhouse experiment to examine the effect of self- and cross-pollination in Mimulus guttatus (Scrophulariaceae) on resistance to and host plant quality for the xylem-feeding spittlebug Philaenus spumarius (Homoptera: Cercopidae). Spittlebugs were found to have a negative effect on two important fitness components in M. guttatus, flower production and above ground biomass. One of two M. guttatus populations examined showed a significant interaction between the pollination and herbivore treatments. In this case, the detrimental effects of herbivores on biomass and flower production were much more pronounced in inbred (self) plants. The presence of spittlebug nymphs increased inbreeding depression by as much as three times. Pollination treatments also had significant effects on important components of herbivore fitness, but these effects were in opposite directions in our two host plant populations. Spittlebug nymphs maturing on self plants emerged as significantly larger adults in one of our host plant populations, indicating that inbreeding increased host plant quality. In our second host plant population, spittlebug nymphs took significantly longer to develop to adulthood on self plants, indicating that inbreeding decreased host plant quality. Taken together these results suggest that the degree of inbreeding in host plant populations can have important and perhaps complex effects on the dynamics of plant-herbivore interactions and on mating-system evolution in the host.     

The role of inbreeding and outbreeding in herbivore resistance and tolerance in Vincetoxicum hirundinaria

Background and Aims

Inbreeding via self-fertilization may have negative effects on plant fitness (i.e. inbreeding depression). Outbreeding, or cross-fertilization between genetically dissimilar parental plants, may also disrupt local adaptation or allelic co-adaptation in the offspring and again lead to reduced plant fitness (i.e. outbreeding depression). Inbreeding and outbreeding may also increase plant vulnerability to natural enemies by altering plant quality or defence. The effects of inbreeding and outbreeding on plant size and response to herbivory in the perennial herb, Vincetoxicum hirundinaria, were investigated.

Methods

Greenhouse experiments were conducted using inbred and outbred (within- and between-population) offspring of 20 maternal plants from four different populations, quantifying plant germination, size, resistance against the specialist folivore, Abrostola asclepiadis, and tolerance of simulated defoliation.

Key Results

Selfed plants were smaller and more susceptible to damage by A. asclepiadis than outcrossed plants. However, herbivore biomass on selfed and outcrossed plants did not differ. The effects of inbreeding on plant performance and resistance did not differ among plant populations or families, and no inbreeding depression at all was found in tolerance of defoliation. Between-population outcrossing had no effect on plant performance or resistance against A. asclepiadis, indicating a lack of outbreeding depression.

Conclusions

Since inbreeding depression negatively affects plant size and herbivore resistance, inbreeding may modify the evolution of the interaction between V. hirundinaria and its specialist folivore. The results further suggest that herbivory may contribute to the maintenance of a mixed mating system of the host plants by selecting for outcrossing and reduced susceptibility to herbivore attack, and thus add to the growing body of evidence on the effects of inbreeding on the mating system evolution of the host plants and the dynamics of plant–herbivore interactions.     

Simultaneous inbreeding modifies inbreeding depression in a plant–herbivore interaction

Because inbreeding is common in natural populations of plants and their herbivores, herbivore‐induced selection on plants, and vice versa, may be significantly modified by inbreeding and inbreeding depression. In a feeding assay with inbred and outbred lines of both the perennial herb, Vincetoxicum hirundinaria, and its specialist herbivore, Abrostola asclepiadis, we discovered that plant inbreeding increased inbreeding depression in herbivore performance in some populations. The effect of inbreeding on plant resistance varied among plant and herbivore populations. The among‐population variation is likely to be driven by variation in plant secondary compounds across populations. In addition, inbreeding depression in plant resistance was substantial when herbivores were outbred, but diminished when herbivores were inbred. These findings demonstrate that in plant–herbivore interactions expression of inbreeding depression can depend on the level of inbreeding of the interacting species. Furthermore, our results suggest that when herbivores are inbred, herbivore‐induced selection against self‐fertilisation in plants may diminish.     

Admixture between native and invasive populations may increase invasiveness of Mimulus guttatus

Self-fertilization and admixture of genotypes from different populations can have major fitness consequences in native species. However, few studies have addressed their potential roles in invasive species. Here, we used plants of Mimulus guttatus from seven native North American, three invasive Scottish and four invasive New Zealand populations to address this. We created seeds from self-fertilization, within-population outcrossing, between-population outcrossing within the same range, and outcrossing between the native and invasive ranges. A greenhouse experiment showed that native and invasive plants of M. guttatus suffered to similar degrees from inbreeding depression, in terms of asexual reproduction and biomass production. After outcrossing with plants from other populations, M. guttatus benefited from heterosis, in terms of asexual and sexual reproduction, and biomass production, particularly when plants from native and invasive populations were crossed. This suggests that, when novel genotypes of M. guttatus from the native North American range will be introduced to the invasive ranges, subsequent outcrossing with M. guttatus plants that are already there might further boost invasiveness of this species.     

Anther-smut infection of Silene alba caused by Ustilago violacea: factors determining fungal reproduction

Summary The anther-smut fungus Ustilago violacea sporulates in flowers of the dioecious host plant Silene alba. Growth chamber comparisons of healthy and diseased plants, with the genetic background of host and pathogen controlled, revealed that fungal infection increases the number of flowers produced per plant and decreases the size of individual flowers. There were few consistent effects of plant genotype or fungal isolate on diseased flower traits, but differences between the plant sexes were apparent. Stimulation of flower production is proportionally greater in females than males: thus, although healthy male plants produce many more flowers than healthy females, sexual differences in diseased flower number are reduced. Sexual differences in diseased flower size also exist, with male flowers smaller than females. A field inoculation study confirmed dimorphism in diseased flower size and demonstrated that spore production per flower was greater for males than females for all flower size classes.     

Little evidence for negative effects of an invasive alien plant on pollinator services

Many invasive alien plants occur in large populations with abundant flowers which are highly attractive to pollinators, and thus might affect pollination of co-occurring native species. This study focuses on the invasive Heracleum mantegazzianum and distance-dependent effects on pollination of Mimulus guttatus in abandoned grassland over 2 years. First, we examined pollinator abundance in yellow traps at 0, 10, 30 and 60–200 m from H. mantegazzianum. We then placed M. guttatus plants at the same distances to monitor effects of the invasive species on pollinator visitation and seed set of neighbouring plants. Finally, we conducted a garden experiment to test if deposition of H. mantegazzianum pollen reduces seed set in M. guttatus. No distance effect was found for the number of bumblebees in traps, although the invasive species attracted a diverse assemblage of insects, and visitation of M. guttatus was enhanced close to H. mantegazzianum. This positive effect was not reflected by seed set of M. guttatus, and heterospecific pollen decreased seed set in these plants. Overall there is little evidence for negative effects of the invasive species on pollination of neighbouring plants, and flower visitation even increases close to the invaded patches. The functional role of the invader and suitable control strategies need further clarification, since removal of H. mantegazzianum may actually damage local pollinator populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Variation in the response of <Emphasis Type="Italic">Mimulus guttatus</Emphasis> (Scrophulariaceae) to herbivore and virus attack

Plants are often simultaneously attacked by insect herbivores and plant pathogens, yet relatively few studies have investigated the potential interactive effects of herbivores and pathogens on plant fitness. We studied the effects of simultaneous attack by meadow spittlebugs, Philaenus spumarius (Homoptera: Cercopidae), and a plant virus, cucumber mosaic virus (CMV), on the fitness of Mimulus guttatus (Scrophulariaceae). We wanted to determine: (1) if trade-offs in defense against meadow spittlebugs and CMV exist, (2) if meadow spittlebugs and CMV interact to affect plant fitness, and (3) if genetic variation is associated with these interactive effects. We found no evidence of trade-offs in defense against meadow spittlebugs and CMV in M. guttatus in a greenhouse experiment. Thus, the ability of M. guttatus to defend itself against one of these enemies is unlikely to preclude the evolution of adequate defenses against the second enemy. We did, however, find strong evidence that spittlebugs and CMV interacted to affect plant fitness and that genetic variation underlies the degree and direction of this interaction. This suggests that selection can act on the genetic variation underlying the interaction between the two enemies and that strong selection imposed by one will alter the response of M. guttatus populations to the second. To our knowledge this is the first study to demonstrate genetic variation associated with the non-additive effect of an herbivore and a pathogen on plant fitness. We suggest that future studies of the mechanisms underlying the defensive properties of plants need to consider variation associated with defense mechanisms and the potential effect of this variation on the response of plant populations to selection by multiple enemies.Co-ordinating editor: Steufer     

The effects of inbreeding, genetic dissimilarity and phenotype on male reproductive success in a dioecious plant

Pollen fate can strongly affect the genetic structure of populations with restricted gene flow and significant inbreeding risk. We established an experimental population of inbred and outbred Silene latifolia plants to evaluate the effects of (i) inbreeding depression, (ii) phenotypic variation and (iii) relatedness between mates on male fitness under natural pollination. Paternity analysis revealed that outbred males sired significantly more offspring than inbred males. Independently of the effects of inbreeding, male fitness depended on several male traits, including a sexually dimorphic (flower number) and a gametophytic trait (in vitro pollen germination rate). In addition, full-sib matings were less frequent than randomly expected. Thus, inbreeding, phenotype and genetic dissimilarity simultaneously affect male fitness in this animal-pollinated plant. While inbreeding depression might threaten population persistence, the deficiency of effective matings between sibs and the higher fitness of outbred males will reduce its occurrence and counter genetic erosion.     

Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems

We examined the effect of self- and cross-pollination on germination success, flowering probability, pollen and ovule production, survivorship, and adult aboveground biomass in two species of Mimulus with contrasting mating systems: the highly seifing M. micranthus and an outcrossing population of M. guttatus. Cross-pollinations were performed both within and between populations in order to examine the scale at which the genetic load is distributed. We found significant inbreeding depression in M. guttatus in four of the six traits, with the highest inbreeding depression observed in biomass (68% and 69% based on within- and between-population crosses, respectively) and lowest in ovule production (21% based on between-population crosses only). M. micranthus displayed significant inbreeding depression in only two of the six traits examined. Again, we observed the highest inbreeding depression in biomass (47–60% based on within- and between-population crosses, respectively), but both traits showing significant differences between self and outcross progeny expressed lower inbreeding depression than in M. guttatus. We detected no significant inbreeding depression for either pollen or ovule production in M. micranthus. An estimate of total inbreeding depression based on the multiplicative effects of all traits was also lower in M. micanthus than∗∗∗ in M. guttatus. Our results are consistent with the expected purging of genetic load in populations with high selfing rates. The absence of inbreeding depression in M. micranthus pollen and ovule production, two traits with strong links to fitness in a selfing annual, further suggests the important role of directional selection in determining the population's genetic load. Comparison of cross-pollinations made within and between populations revealed little evidence of divergence of genetic load among the M. micranthus and M. guttatus populations examined.     

REPRODUCTIVE ASSURANCE MECHANISMS IN THREE TAXA OF THE MIMULUS GUTTATUS COMPLEX (SCROPHULARIACEAE)

The evolution of inbreeding in plants has often been attributed to selection for the ability to set seed in the absence of mates or pollinators. Mechanisms of reproductive assurance in five populations of mixed mating Mimulus guttatus, three populations of inbreeding M. platycalyx, and two populations of inbreeding M. nasutus were examined in a pollinator-free greenhouse. Reproductive assurance was manifested in all populations by autofertility, vegetative reproduction, or both. The inbreeding taxa had significantly greater levels of autofertility and less vegetative reproduction. Three modes of autofertility were identified: 1) due to corolla abscission only, occurring in three M. guttatus populations; 2) due to both corolla abscission and direct anther-stigma contact by curling of the lower stigmatic lobe into the anthers, occurring in two M. guttatus populations; and 3) direct stigma-anther contact by stigma curling alone prior to corolla abscission, found in each M. platycalyx and M. nasutus population. Stigma-anther distance and its interaction with stigma curling contributed to differences in autofertility among populations. Significant levels of intrapopulation quantitative genetic variation were found for seven of ten traits examined; average levels were similar between inbreeding and mixed mating populations. Genetic variation within populations for autofertility per se was not detected, but significant levels controlling stigma-anther distance were found in two M. guttatus populations. These results show that evolution of inbreeding by natural selection for reproductive assurance is possible in Mimulus, and illustrate the complex changes in floral dynamics and morphology it may involve.     

Preference for outbred host plants and positive effects of inbreeding on egg survival in a specialist herbivore

Inbreeding can profoundly affect the interactions of plants with herbivores as well as with the natural enemies of the herbivores. We studied how plant inbreeding affects herbivore oviposition preference, and whether inbreeding of both plants and herbivores alters the probability of predation or parasitism of herbivore eggs. In a laboratory preference test with the specialist herbivore moth Abrostola asclepiadis and inbred and outbred Vincetoxicum hirundinaria plants, we discovered that herbivores preferred to oviposit on outbred plants. A field experiment with inbred and outbred plants that bore inbred or outbred herbivore eggs revealed that the eggs of the outbred herbivores were more likely to be lost by predation, parasitism or plant hypersensitive responses than inbred eggs. This difference did not lead to differences in the realized fecundity as the number of hatched larvae did not differ between inbred and outbred herbivores. Thus, the strength of inbreeding depression in herbivores decreases when their natural enemies are involved. Plant inbreeding did not alter the attraction of natural enemies of the eggs. We conclude that inbreeding can significantly alter the interactions of plants and herbivores at different life-history stages, and that some of these alterations are mediated by the natural enemies of the herbivores.     

Inbreeding and extinction: Effects of purging

Deleterious alleles may be removed (purged) bynatural selection in populations undergoinginbreeding. However, there is controversyregarding the effectiveness of purging inreducing the extinction risk due to inbreeding,particularly in conservation contexts. Weevaluated the effects of purging on theextinction risk due to inbreeding in Drosophila melanogaster using two basepopulations, an outbred population (non-purged)and four-way crosses between highly inbredlines derived from the same population(purged). The inbred lines used in the four-waycrosses were previously subjected to 20generations of full-sib mating. The impact offull-sib inbreeding over a further 12generations was compared in 200 populationsfrom each of the two base populations. Therewas a small and non-significant differencebetween the extinction rates at an inbreedingcoefficient of 0.93 in the non-purged (0.74± 0.03) and purged (0.69 ± 0.03)treatments. This is consistent with otherevidence indicating that the effects of purgingare often small. Purging using rapid inbreedingin very small populations cannot be relied uponto eliminate the deleterious effects ofinbreeding.     

Variation in life-history traits among Urtica dioica populations with different history in parasitism by the holoparasitic plant Cuscuta europaea

Several studies demonstrate that natural enemies (e.g. parasites) have profound negative effects on the life-history traits of their hosts. If the host can compensate for the negative effects of parasitic infection by altering its life history, these modifications may partly form the basis of resistance or tolerance against parasites. Thus, parasites may be of considerable importance in shaping the evolution of life-history traits of their hosts. To examine if previous parasitism is associated with differences in life-history traits of the host, I conducted a common garden experiment with Urtica dioica plants originating from eight populations of which four were unparasitized, and four parasitized by the holoparasitic plant, Cuscuta europaea. A field survey indicated no differences between unparasitized and parasitized populations in, for example, the number of plant species and nutrient levels in the soil. Thus, it seems reasonable to assume that differences in life-history traits between the two population types in the common garden would reflect the effects of previous selection by the parasite. In the common garden, plants from parasitized populations started to flower later and allocated less biomass to asexual reproduction (measured as the production of stolons, i.e. clonal propagation) compared to plants from unparasitized populations. These results thus indicate that selection by the parasite may have favoured later onset of flowering, and may have selected against asexual reproduction.     

MEASURES OF PHENOTYPIC SELECTION ARE BIASED BY PARTIAL INBREEDING

When populations are partially inbred due to the population structure or to a mixed mating system like partial self-fertilization, some individuals will be more inbred than others. This heterogeneity among individuals in the history of inbreeding can greatly complicate the interpretation of measures of quantitative genetic variability when the traits studied exhibit inbreeding depression. Partial inbreeding can also bias measures of phenotypic selection toward the detection of strong directional and stabilizing selection. In this paper, data are presented from several inbreeding experiments conducted on two partially selfing, annual populations of the monkeyflower Mimulus guttatus that show that the means of many of the morphological and phenological traits measured were affected by inbreeding. These findings imply that estimates of heritabilities and additive genetic covariances would not reflect the potential for these populations to respond to selection. Phenotypic selection analyses conducted on naturally occurring plants, involving linear regressions of relative seed production on the traits, revealed significant directional selection on many of the same quantitative traits measured in the inbreeding studies. However, when the same selection analyses were performed on plants with known histories of inbreeding, part of the statistical relationship between relative seed number and the traits was found to be due to the mating system: inbred individuals had both lower seed production and different mean values for the traits than outcrossed individuals. It is also shown, with a hypothetical example, that partial inbreeding can bias measures of stabilizing selection toward the detection of strong stabilizing selection. Partial inbreeding therefore tends to make directional and stabilizing selection appear stronger than it is, and it may be that natural selection in the wild is actually weaker than many studies of partially inbred species suggest.     

The effect of inbreeding on defence against multiple enemies in Datura stramonium

The ability of plants to respond to natural enemies might depend on the availability of genetic variation for the optimal phenotypic expression of defence. Selfing can affect the distribution of genetic variability of plant fitness, resistance and tolerance to herbivores and pathogens. The hypothesis of inbreeding depression influencing plant defence predicts that inbreeding would reduce resistance and tolerance to damage by natural enemies relative to outcrossing. In a field experiment entailing experimentally produced inbred and outcrossed progenies, we assessed the effects of one generation of selfing on Datura stramonium resistance and tolerance to three types of natural enemies, herbivores, weevils and a virus. We also examined the effect of damage on relative growth rate (RGR), flower, fruit, and seed production in inbred and outcrossed plants. Inbreeding significantly reduced plant defence to natural enemies with an increase of 4% in herbivore damage and 8% in viral infection. These results indicate inbreeding depression in total resistance. Herbivory increased 10% inbreeding depression in seed number, but viral damage caused inbred and outcrossed plants to have similar seed production. Inbreeding and outcrossing effects on fitness components were highly variable among families, implying that different types or numbers of recessive deleterious alleles segregate following inbreeding in D. stramonium. Although inbreeding did not equally alter all the interactions, our findings indicate that inbreeding reduced plant defence to herbivores and pathogens in D. stramonium.     

Local pre‐adaptation to disturbance and inbreeding–environment interactions affect colonisation abilities of diploid and tetraploid Centaurea stoebe

• Primary colonisation in invasive ranges most commonly occurs in disturbed habitats, where anthropogenic disturbance may cause physical damage to plants. The tolerance to such damage may differ between cytotypes and among populations as a result of differing population histories (adaptive differentiation between ruderal verus natural habitats). Moreover, founder populations often experience inbreeding depression, the effects of which may increase through physical damage due to inbreeding–environment interactions. We aimed to understand how such colonisation processes differ between diploid and tetraploid Centaurea stoebe populations, with a view to understanding why only tetraploids are invasive.
• We conducted a clipping experiment (frequency: zero, once or twice in the growing season) on inbred versus outbred offspring originating from 37 C. stoebe populations of varying cytotype, range and habitat type (natural versus ruderal). Aboveground biomass was harvested at the end of the vegetation period, while re‐sprouting success was recorded in the following spring.
• Clipping reduced re‐sprouting success and biomass, which was significantly more pronounced in natural than in ruderal populations. Inbreeding depression was not detected under benign conditions, but became increasingly apparent in biomass when plants were clipped. The effects of clipping and inbreeding did not differ between cytotypes.
• Adaptive differentiation in disturbance tolerance was higher among populations than between cytotypes, which highlights the potential of pre‐adaptation in ruderal populations during early colonisation on anthropogenically disturbed sites. While the consequences of inbreeding increased through clipping‐mediated stress, they were comparable between cytotypes, and consequently do not contribute to understanding the cytotype shift in the invasive range.

     

Chloroplast and nuclear DNA variation among homozygous plants in a population of the autogamous annualMicroseris douglasii (Asteraceae,Lactuceae)

The autogamous diploid annualMicroseris douglasii of California occurs in many isolated populations. The populations consist of one to many highly inbred biotypes. Morphological variation among populations usually is greater than within populations. In spite of the virtual absence of gene flow even within populations, genetically determined character differences are randomly distributed and associated throughout the range of the species. Recent evidence even suggests introgression of chloroplasts from the relatedM. bigelovii. Offspring families from 25 plants of a very variable population were raised and examined for segregation of morphological and molecular (RAPD) markers. All 25 original plants were completely homozygous for all markers, but each differed from all others at least in some markers. The population consisted of two genetically isolated groups of plants: a distinct inbred line (3 plants) and 22 plants with random associations of a common set of markers and characters, possibly recombinant inbreds from a past hybridization event. One of these 22 plants contained a chloroplast genome found inM. bigelovii, the other 24 plants a chloroplast genome found only inM. douglasii.