Tests for the joint evolution of mating system and drought escape in Mimulus

Background and Aims

Self-fertilizing taxa are often found at the range margins of their progenitors, where sub-optimal habitats may select for alternative physiological strategies. The extent to which self-fertilization is favoured directly vs. arising indirectly through correlations with other adaptive life history traits is unclear. Trait responses to selection depend on genetic variation and covariation, as well as phenotypic and genetic responses to altered environmental conditions. We tested predictions of the hypothesis that self-fertilization in Mimulus arises through direct selection on physiological and developmental traits that allow seasonal drought escape.

Methods

Phenotypic selection on mating system and drought escape traits was estimated in field populations of M. guttatus. In addition, trait phenotype and phenotypic selection were compared between experimental wet and dry soil in two greenhouse populations each of M. guttatus and M. nasutus. Finally, genetic variation and covariation for traits were compared between wet and dry soil treatments in a greenhouse population of M. guttatus.

Key Results

Consistent with predictions, selection for early flowering was generally stronger than for mating system traits, and selection for early flowering was stronger in dry soil. Inconsistent with predictions, selection for water-use efficiency was largely absent; selection for large flowers was stronger than for drought escape in the field; and most drought escape and mating system traits were not genetically correlated. A positive genetic correlation between flowering time and flower size, which opposed the adaptive contour, emerged only in wet soil, suggesting that variation in water availability may maintain variation in these traits. Plastic responses to soil moisture treatments supported the idea that taxonomic divergence could have been facilitated by plasticity in flowering time and selfing.

Conclusions

The hypothesis that plant mating systems may evolve indirectly via selection on correlated life history characteristics is plausible and warrants increased attention.     

Plant resource-use strategies: the importance of phenotypic plasticity in response to a productivity gradient for two subalpine species

Background and Aims

Functional traits are indicators of plant interactions with their environment and the resource-use strategies of species can be defined through some key functional traits. The importance of genetic variability and phenotypic plasticity in trait variations in response to a common environmental change was investigated in two subalpine species.

Methods

Two species with contrasted resource-use strategies, Dactylis glomerata and Festuca paniculata, were grown along a productivity gradient in a greenhouse experiment. Functional traits of different genotypes were measured to estimate the relative roles of phenotypic plasticity and genetic variability, and to compare their levels of phenotypic plasticity.

Key Results

Trait variability in the field for the two species is more likely to be the result of phenotypic plasticity rather than of genetic differentiation between populations. The exploitative species D. glomerata expressed an overall higher level of phenotypic plasticity compared with the conservative species F. paniculata. In addition to different amplitudes of phenotypic plasticity, the two species differed in their pattern of response for three functional traits relevant to resource use (specific leaf area, leaf dry matter content and leaf nitrogen content).

Conclusions

Functional trait variability was mainly the result of phenotypic plasticity, with the exploitative species showing greater variability. In addition to average trait values, two species with different resource-use strategies differed in their plastic responses to productivity.     

Molecular and quantitative trait variation within and among small fragmented populations of the endangered plant species Psilopeganum sinense

Background and Aims

Natural selection and genetic drift are important evolutionary forces in determining genetic and phenotypic differentiation in plant populations. The extent to which these two distinct evolutionary forces affect locally adaptive quantitative traits has been well studied in common plant and animal species. However, we know less about how quantitative traits respond to selection pressures and drift in endangered species that have small population sizes and fragmented distributions. To address this question, this study assessed the relative strengths of selection and genetic drift in shaping population differentiation of phenotypic traits in Psilopeganum sinense, a naturally rare and recently endangered plant species.

Methods

Population differentiation at five quantitative traits (Q_ST) obtained from a common garden experiment was compared with differentiation at putatively neutral microsatellite markers (F_ST) in seven populations of P. sinense. Q_ST estimates were derived using a Bayesian hierarchical variance component method.

Key Results

Trait-specific Q_ST values were equal to or lower than F_ST. Neutral genetic diversity was not correlated with quantitative genetic variation within the populations of P. sinense.

Conclusions

Despite the prevalent empirical evidence for Q_ST > F_ST, the results instead suggest a definitive role of stabilizing selection and drift leading to phenotypic differentiation among small populations. Three traits exhibited a significantly lower Q_ST relative to F_ST, suggesting that populations of P. sinense might have experienced stabilizing selection for the same optimal phenotypes despite large geographical distances between populations and habitat fragmentation. For the other two traits, Q_ST estimates were of the same magnitude as F_ST, indicating that divergence in these traits could have been achieved by genetic drift alone. The lack of correlation between molecular marker and quantitative genetic variation suggests that sophisticated considerations are required for the inference of conservation measures of P. sinense from neutral genetic markers.     

Divergent selection in trailing- versus leading-edge populations of Biscutella laevigata

Background and Aims

Knowledge on how climate-induced range shifts might affect natural selection is crucial to understand the evolution of species ranges.

Methods

Using historical demographic perspectives gathered from regional-scale phylogeography on the alpine herb Biscutella laevigata, indirect inferences on gene flow and signature of selection based on AFLP genotyping were compared between local populations persisting at the trailing edge and expanding at the leading edge.

Key Results

Spatial autocorrelation revealed that gene flow was two times more restricted at the trailing edge and genome scans indicated divergent selection in this persisting population. In contrast, no pattern of selection emerged in the expanding population at the leading edge.

Conclusions

Historical effects may determine different architecture of genetic variation and selective patterns within local populations, what is arguably important to understand evolutionary processes acting across the species ranges.     

Intraspecific variation in the magnitude and pattern of flooding-induced shoot elongation in Rumex palustris

Background and Aims

Intraspecific variation in flooding tolerance is the basic pre-condition for adaptive flooding tolerance to evolve, and flooding-induced shoot elongation is an important trait that enables plants to survive shallow, prolonged flooding. Here an investigation was conducted to determine to what extent variation in flooding-induced leaf elongation exists among and within populations of the wetland species Rumex palustris, and whether the magnitude of elongation can be linked to habitat characteristics.

Methods

Offspring of eight genotypes collected in each of 12 populations from different sites (ranging from river mudflats with dynamic flooding regimes to areas with stagnant water) were submerged, and petioles, laminas and roots were harvested separately to measure traits related to elongation and plant growth.

Key Results

We found strong elongation of petioles upon submergence, and both among- and within-population variation in this trait, not only in final length, but also in the timing of the elongation response. However, the variation in elongation responses could not be linked to habitat type.

Conclusions

Spatio-temporal variation in the duration and depth of flooding in combination with a presumably weak selection against flooding-induced elongation may have contributed to the maintenance of large genetic variation in flooding-related traits among and within populations.     

Exogenous selection shapes germination behaviour and seedling traits of populations at different altitudes in a Senecio hybrid zone

Background and Aims

The Senecio hybrid zone on Mt Etna, Sicily, is characterized by steep altitudinal clines in quantitative traits and genetic variation. Such clines are thought to be maintained by a combination of ‘endogenous’ selection arising from genetic incompatibilities and environment-dependent ‘exogenous’ selection leading to local adaptation. Here, the hypothesis was tested that local adaptation to the altitudinal temperature gradient contributes to maintaining divergence between the parental species, S. chrysanthemifolius and S. aethnensis.

Methods

Intra- and inter-population crosses were performed between five populations from across the hybrid zone and the germination and early seedling growth of the progeny were assessed.

Key Results

Seedlings from higher-altitude populations germinated better under low temperatures (9–13 °C) than those from lower altitude populations. Seedlings from higher-altitude populations had lower survival rates under warm conditions (25/15 °C) than those from lower altitude populations, but also attained greater biomass. There was no altitudinal variation in growth or survival under cold conditions (15/5 °C). Population-level plasticity increased with altitude. Germination, growth and survival of natural hybrids and experimentally generated F₁s generally exceeded the worse-performing parent.

Conclusions

Limited evidence was found for endogenous selection against hybrids but relatively clear evidence was found for divergence in seed and seedling traits, which is probably adaptive. The combination of low-temperature germination and faster growth in warm conditions might enable high-altitude S. aethnensis to maximize its growth during a shorter growing season, while the slower growth of S. chrysanthemifolius may be an adaptation to drought stress at low altitudes. This study indicates that temperature gradients are likely to be an important environmental factor generating and maintaining adaptive divergence across the Senecio hybrid zone on Mt Etna.     

Evolution of pollination niches and floral divergence in the generalist plant Erysimum mediohispanicum

Background and Aims

How generalist plants diverge in response to pollinator selection without becoming specialized is still unknown. This study explores this question, focusing on the evolution of the pollination system in the pollination generalist Erysimum mediohispanicum (Brassicaceae).

Methods

Pollinator assemblages were surveyed from 2001 to 2010 in 48 geo-referenced populations covering the entire geographic distribution of E. mediohispanicum. Bipartite modularity, a complex network tool, was used to find the pollination niche of each population. Evolution of the pollination niches and the correlated evolution of floral traits and pollination niches were explored using within-species comparative analyses.

Key Results

Despite being generalists, the E. mediohispanicum populations studied can be classified into five pollination niches. The boundaries between niches were not sharp, the niches differing among them in the relative frequencies of the floral visitor functional groups. The absence of spatial autocorrelation and phylogenetic signal indicates that the niches were distributed in a phylogeographic mosaic. The ancestral E. mediohispanicum populations presumably belonged to the niche defined by a high number of beetle and ant visits. A correlated evolution was found between pollination niches and some floral traits, suggesting the existence of generalist pollination ecotypes.

Conclusions

It is conjectured that the geographic variation in pollination niches has contributed to the observed floral divergence in E. mediohispanicum. The process mediating this floral divergence presumably has been adaptive wandering, but the adaptation to the local pollinator faunas has been not universal. The outcome is a landscape where a few populations locally adapted to their pollination environment (generalist pollination ecotypes) coexist with many populations where this local adaptation has failed and where the plant phenotype is not primarily shaped by pollinators.     

Variation in inbreeding depression and plasticity across native and non-native field environments

Background and Aims

Since the early 1990s, research on genetic variation of phenotypic plasticity has expanded and empirical research has emphasized the role of the environment on the expression of inbreeding depression. An emerging question is how these two evolutionary ecology mechanisms interact in novel environments. Interest in this area has grown with the need to understand the establishment of populations in response to climate change, and to human-assisted transport to novel environments.

Methods

We compare performance in the field of outcrossed (O) and inbred lines (S1, S2) from 20 maternal families from each of two native populations of Mimulus guttatus. The experiment was planted in California in each population''s home site, in the other populations''s home site, in a novel site within the native range of M. guttatus, and in a novel site within the non-native range in North America. The experiment included nearly 6500 individuals. Survival, sexual reproduction and above-ground biomass were examined in order to evaluate inbreeding depression, and stem diameter and plant height were examined in order to evaluate phenotypic plasticity.

Key Results

Across all field sites, approx. 36 % of plants survived to flowering. Inbreeding depression differed among sites and outcrossed offspring generally outperformed selfed offspring. However, in the native-novel site, self-progeny performed better or equally well as outcross progeny. Significant phenotypic plasticity and genetic variation in plasticity was detected in the two architectural traits measured. The absolute value of plasticity showed the most marked difference between home and non-native novel site or non-native-novel site. Evidence was detected for an interaction between inbreeding and plasticity for stem diameter.

Conclusions

The results demonstrate that during initial population establishment, both inbreeding depression and phenotypic plasticity vary among field sites, and may be an important response to environments outside a species'' currently occupied range. However, the interaction between inbreeding and plasticity may be limited and environment-dependent.     

Ontogenetic contingency of tolerance mechanisms in response to apical damage

Background and Aims

Plants are able to tolerate tissue loss through vigorous branching which is often triggered by release from apical dominance and activation of lateral meristems. However, damage-induced branching might not be a mere physiological outcome of released apical dominance, but an adaptive response to environmental signals, such as damage timing and intensity. Here, branching responses to both factors were examined in the annual plant Medicago truncatula.

Methods

Branching patterns and allocation to reproductive traits were examined in response to variable clipping intensities and timings in M. truncatula plants from two populations that vary in the onset of reproduction. Phenotypic selection analysis was used to evaluate the strength and direction of selection on branching under the damage treatments.

Key Results

Plants of both populations exhibited an ontogenetic shift in tolerance mechanisms: while early damage induced greater meristem activation, late damage elicited investment in late-determined traits, including mean pod and seed biomass, and supported greater germination rates. Severe damage mostly elicited simultaneous development of multiple-order lateral branches, but this response was limited to early damage. Selection analyses revealed positive directional selection on branching in plants under early- compared with late- or no-damage treatments.

Conclusions

The results demonstrate that damage-induced meristem activation is an adaptive response that could be modified according to the plant''s developmental stage, severity of tissue loss and their interaction, stressing the importance of considering these effects when studying plastic responses to apical damage.     

Polyploidy influences sexual system and mating patterns in the moss Atrichum undulatum sensu lato

Background and Aims

Evolutionary transitions between separate and combined sexes have frequently occurred across various plant lineages. In mosses, which are haploid-dominant, evolutionary transitions from separate to combined sexes are often associated with genome doubling. Polyploidy and hermaphroditism have strong effects on the inbreeding depression of a population, and are subsequently predicted to affect the mating system.

Methods

We tested the association between ploidy (haploid, diploid or triploid gametophytes) and mating system in 21 populations of Atrichum undulatum sensu lato, where sex ratios vary widely. For each population, we measured the sex ratio, estimated selfing rates using allozyme markers and determined the level of ploidy through flow cytometry.

Key Results

Hermaphrodites in A. undulatum were either diploid or triploid. However, many diploid populations were strictly separate-sexed, suggesting that hermaphroditism is not a necessary result of genome doubling. Levels of selfing were strongly supported as being greater than zero in one population with strictly separate-sexed individuals, and one-third of populations with hermaphrodites.

Conclusions

Although hermaphrodites are associated with triploidy, hermaphroditism is not a necessary outcome of genome duplication. Hermaphroditism, but not genome duplication alone, increased estimated selfing rates, probably due to the occurrence of selfing within a gametophyte. Thus, genome duplication can influence the mating system and the associated evolution and maintenance of reproductive traits.     

Novel adaptation to hawkmoth pollinators in Clarkia reduces efficiency,not attraction of diurnal visitors

Background and Aims

Plant populations experiencing divergent pollination environments may be under selection to modify floral traits in ways that increase both attractiveness to and efficiency of novel pollinators. These changes may come at the cost of reducing overall effectiveness of other pollinators. The goal of this study was to examine differences in attractiveness and efficiency between Clarkia concinna and C. breweri, sister species of annual plants with parapatric distributions.

Methods

An assessment was made as to whether observed differences in visitors between natural populations are driven by differences in floral traits or differences in the local pollination environment. Differences in floral attractiveness were quantified by setting out arrays of both species in the geographical range of each species and exposing both species to nocturnal hawkmoths (Hyles lineata) in flight cages. Differences in visitor efficiency were estimated by measuring stigma–visitor contact frequency and pollen loads for diurnal visitors, and pollen deposition on stigmas for hawkmoths.

Key Results

The composition of visitors to arrayed plants was similar between plant species at any particular site, but highly divergent among sites, and reflected differences in visitors to natural populations. Diurnal insects visited both species, but were more common at C. concinna populations. Hummingbirds and hawkmoths were only observed visiting within the range of C. breweri. Despite attracting similar species when artificially presented together, C. concinna and C. breweri showed large differences in pollinator efficiency. All visitors except hawkmoths pollinated C. concinna more efficiently.

Conclusions

Differences in the available pollinator community may play a larger role than differences in floral traits in determining visitors to natural populations of C. concinna and C. breweri. However, floral traits mediate differences in pollinator efficiency. Increased effectiveness of the novel hawkmoth pollinator on C. breweri comes at relatively little cost in attractiveness to other visitors, but at large cost in their efficiency as pollinators.     

Experimentally reduced root–microbe interactions reveal limited plasticity in functional root traits in Acer and Quercus

Background and Aims

Interactions between roots and soil microbes are critical components of below-ground ecology. It is essential to quantify the magnitude of root trait variation both among and within species, including variation due to plasticity. In addition to contextualizing the magnitude of plasticity relative to differences between species, studies of plasticity can ascertain if plasticity is predictable and whether an environmental factor elicits changes in traits that are functionally advantageous.

Methods

To compare functional traits and trait plasticities in fine root tissues with natural and reduced levels of colonization by microbial symbionts, trimmed and surface-sterilized root segments of 2-year-old Acer rubrum and Quercus rubra seedlings were manipulated. Segments were then replanted into satellite pots filled with control or heat-treated soil, both originally derived from a natural forest. Mycorrhizal colonization was near zero in roots grown in heat-treated soil; roots grown in control soil matched the higher colonization levels observed in unmanipulated root samples collected from field locations.

Key Results

Between-treatment comparisons revealed negligible plasticity for root diameter, branching intensity and nitrogen concentration across both species. Roots from treated soils had decreased tissue density (approx. 10–20 %) and increased specific root length (approx. 10–30 %). In contrast, species differences were significant and greater than treatment effects in traits other than tissue density. Interspecific trait differences were also significant in field samples, which generally resembled greenhouse samples.

Conclusions

The combination of experimental and field approaches was useful for contextualizing trait plasticity in comparison with inter- and intra-specific trait variation. Findings that root traits are largely species dependent, with the exception of root tissue density, are discussed in the context of current literature on root trait variation, interactions with symbionts and recent progress in standardization of methods for quantifying root traits.     

How slug herbivory of juvenile hybrid willows alters chemistry,growth and subsequent susceptibility to diverse plant enemies

Background and Aims

Selective feeding by herbivores, especially at the seedling or juvenile phase, has the potential to change plant traits and ultimately the susceptibility of surviving plants to other enemies. Moreover, since hybridization is important to speciation and can lead to introgression of traits between plant species, differential feeding (herbivore-induced mortality) can influence the expression of resistance traits of hybrids and ultimately determine the consequences of hybridization. While it would be expected that herbivore-induced mortality would lead to greater resistance, there may be trade-offs whereby resistance to one herbivore increases susceptibility to others. The hypothesis was tested that the exotic slug, Arion subfuscus, causes non-random survival of hybrid willows and alters plant: (1) susceptibility to slugs; (2) secondary and nutritional chemistry, and growth; and (3) susceptibility to other phytophages.

Methods

Two populations of plants, control and selected, were created by placing trays of juvenile willows in the field and allowing slugs access to only some. When ≤10 individuals/tray remained (approx. 85 % mortality), ‘selected’ and undamaged ‘control’ trays were returned to a common area. Traits of these populations were then examined in year 1 and in subsequent years.

Key Results

The selected population was less palatable to slugs. Surprisingly, foliar concentrations of putative defence traits (phenolic glycosides and tannins) did not differ between treatments, but the selected population had higher foliar nitrogen and protein, lower carbon to nitrogen ratio and greater above-ground biomass, indicating that vigorously growing plants were inherently more resistant to slugs. Interestingly, selected plants were more susceptible to three phytophages: an indigenous pathogen (Melampsora epitea), a native herbivorous beetle (Chrysomela knabi) and an exotic willow leaf beetle (Plagiodera versicolora).

Conclusions

This exotic slug changed the population structure of F₂ hybrid willows in unanticipated ways. Defence expression remained unchanged, while nutritional and growth traits changed. These changes caused plants to be more susceptible to other plant enemies. Other exotic herbivore species are anticipated to have similar direct and indirect effects on native plant populations.     

Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome

Background and Aims

If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.

Methods

Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.

Key Results

Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.

Conclusions

This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.     

Gender plasticity and sexual system stability in Wurmbea

Background and aims

Sexually dimorphic populations are often located in drier habitats than cosexual populations. Gender plasticity (GP), whereby hermaphrodites alter female and male functions depending on resources, and sex-differential plasticity (SDP) between hermaphrodites and unisexuals are predicted to affect sexual system stability. Here, GP and SDP are evaluated in cosexual and gynodioecious Wurmbea biglandulosa and sub-dioecious and dioecious W. dioica.

Methods

GP was evaluated under two resource conditions, compared among sexual systems and assessed as to whether (1) males produced perfect flowers and (2) hermaphrodites altered investment in perfect (female function) and total (male function) flowers. SDP was assessed within sexual systems as differences between sex functions of hermaphrodites vs. unisexuals. Males and hermaphrodites were compared to assess whether size thresholds for female function differed among sexual systems. Plasticity costs were evaluated using correlations between female function and male traits in hermaphrodites, and in W. dioica by comparing hermaphrodite and male regressions between plant size and pollen production.

Key Results

In dioecious W. dioica no males exhibited GP, whereas 100 % did in gynodioecious and cosexual W. biglandulosa. In sub-dioecious W. dioica, resources affected GP (high, 66 %; low, 42 %). Hermaphrodites in all sexual systems reduced perfect but not total flowers under low resources. Unisexuals were unaffected, demonstrating SDP for female function only. Thresholds for female function were greater in sub-dioecious W. dioica than in W. biglandulosa. Plasticity costs were detected only in sub-dioecious W. dioica.

Conclusions

SDP for female function could assist female establishment in cosexual populations and maintain females in gynodioecious and sub-dioecious populations. Although the absence of male SDP should stabilize sub-dioecy, plasticity costs would render sub-dioecy unstable, favouring canalized males over hermaphrodites. This study highlights the importance of interactions between environmental conditions and hermaphrodite sex expression for the stability of dimorphic sexual systems.     

Long-term functional plasticity in plant hydraulic architecture in response to supplemental moisture

Background and Aims

Plasticity in structural and functional traits related to water balance may determine plant performance and survival in ecosystems characterized by water limitation or high levels of rainfall variability, particularly in perennial herbaceous species with long generation cycles. This paper addresses whether and the extent to which several such seasonal to long-term traits respond to changes in moisture availability.

Methods

Using a novel approach that integrates ecology, physiology and anatomy, a comparison was made of lifetime functional traits in the root xylem of a long-lived perennial herb (Potentilla diversifolia, Rosaceae) growing in dry habitats with those of nearby individuals growing where soil moisture had been supplemented for 14 years. Traditional parameters such as specific leaf area (SLA) and above-ground growth were also assessed.

Key Results

Individuals from the site receiving supplemental moisture consistently showed significant responses in all considered traits related to water balance: SLA was greater by 24 %; roots developed 19 % less starch storing tissue, an indicator for drought-stress tolerance; and vessel size distributions shifted towards wider elements that collectively conducted water 54 % more efficiently – but only during the years for which moisture was supplemented. In contrast, above-ground growth parameters showed insignificant or inconsistent responses.

Conclusions

The phenotypic changes documented represent consistent, dynamic responses to increased moisture availability that should increase plant competitive ability. The functional plasticity of xylem anatomy quantified in this study constitutes a mechanistic basis for anticipating the differential success of plant species in response to climate variability and change, particularly where water limitation occurs.     

Highly local environmental variability promotes intrapopulation divergence of quantitative traits: an example from tropical rain forest trees

Background and Aims

In habitat mosaics, plant populations face environmental heterogeneity over short geographical distances. Such steep environmental gradients can induce ecological divergence. Lowland rainforests of the Guiana Shield are characterized by sharp, short-distance environmental variations related to topography and soil characteristics (from waterlogged bottomlands on hydromorphic soils to well-drained terra firme on ferralitic soils). Continuous plant populations distributed along such gradients are an interesting system to study intrapopulation divergence at highly local scales. This study tested (1) whether conspecific populations growing in different habitats diverge at functional traits, and (2) whether they diverge in the same way as congeneric species having different habitat preferences.

Methods

Phenotypic differentiation was studied within continuous populations occupying different habitats for two congeneric, sympatric, and ecologically divergent tree species (Eperua falcata and E. grandiflora, Fabaceae). Over 3000 seeds collected from three habitats were germinated and grown in a common garden experiment, and 23 morphological, biomass, resource allocation and physiological traits were measured.

Key Results

In both species, seedling populations native of different habitats displayed phenotypic divergence for several traits (including seedling growth, biomass allocation, leaf chemistry, photosynthesis and carbon isotope composition). This may occur through heritable genetic variation or other maternally inherited effects. For a sub-set of traits, the intraspecific divergence associated with environmental variation coincided with interspecific divergence.

Conclusions

The results indicate that mother trees from different habitats transmit divergent trait values to their progeny, and suggest that local environmental variation selects for different trait optima even at a very local spatial scale. Traits for which differentiation within species follows the same pattern as differentiation between species indicate that the same ecological processes underlie intra- and interspecific variation.     

Genetic evidence for predominantly hydrochoric gene flow in the invasive riparian plant Impatiens glandulifera (Himalayan balsam)

Background and Aims

Riparian systems are prone to invasion by alien plant species. The spread of invasive riparian plants may be facilitated by hydrochory, the transport of seeds by water, but while ecological studies have highlighted the possible role of upstream source populations in the establishment and persistence of stands of invasive riparian plant species, population genetic studies have as yet not fully addressed the potential role of hydrochoric dispersal in such systems.

Methods

A population genetics approach based on a replicated bifurcate sampling design is used to test hypotheses consistent with patterns of unidirectional, linear gene flow expected under hydrochoric dispersal of the invasive riparian plant Impatiens glandulifera in two contrasting river systems.

Key results

A significant increase in levels of genetic diversity downstream was observed, consistent with the accumulation of propagules from upstream source populations, and strong evidence was found for organization of this diversity between different tributaries, reflecting the dendritic organization of the river systems studied.

Conclusions

These findings indicate that hydrochory, rather than anthropogenic dispersal, is primarily responsible for the spread of I. glandulifera in these river systems, and this is relevant to potential approaches to the control of invasive riparian plant species.     

Invasion strategies in clonal aquatic plants: are phenotypic differences caused by phenotypic plasticity or local adaptation?

Background and Aims

The successful spread of invasive plants in new environments is often linked to multiple introductions and a diverse gene pool that facilitates local adaptation to variable environmental conditions. For clonal plants, however, phenotypic plasticity may be equally important. Here the primary adaptive strategy in three non-native, clonally reproducing macrophytes (Egeria densa, Elodea canadensis and Lagarosiphon major) in New Zealand freshwaters were examined and an attempt was made to link observed differences in plant morphology to local variation in habitat conditions.

Methods

Field populations with a large phenotypic variety were sampled in a range of lakes and streams with different chemical and physical properties. The phenotypic plasticity of the species before and after cultivation was studied in a common garden growth experiment, and the genetic diversity of these same populations was also quantified.

Key Results

For all three species, greater variation in plant characteristics was found before they were grown in standardized conditions. Moreover, field populations displayed remarkably little genetic variation and there was little interaction between habitat conditions and plant morphological characteristics.

Conclusions

The results indicate that at the current stage of spread into New Zealand, the primary adaptive strategy of these three invasive macrophytes is phenotypic plasticity. However, while limited, the possibility that genetic diversity between populations may facilitate ecotypic differentiation in the future cannot be excluded. These results thus indicate that invasive clonal aquatic plants adapt to new introduced areas by phenotypic plasticity. Inorganic carbon, nitrogen and phosphorous were important in controlling plant size of E. canadensis and L. major, but no other relationships between plant characteristics and habitat conditions were apparent. This implies that within-species differences in plant size can be explained by local nutrient conditions. All together this strongly suggests that invasive clonal aquatic plants adapt to a wide range of habitats in introduced areas by phenotypic plasticity rather than local adaptation.