Experimental life-history evolution: selection on the allocation to sexual reproduction and its plasticity in a clonal plant

Abstract.— Allocation to sexual reproduction is an important life-history trait in clonal plants. Different selection pressures between competitive and competition-free environments are likely to result in the evolution of specialized genotypes and to maintain genetic variation in reproductive allocation. Moreover, selection may also result in the evolution of plastic allocation strategies. The necessary prerequisite for evolution, heritable genetic variation, can best be studied with selection experiments. Starting from a base population of 102 replicated genotypes of the clonal herb Ranunculus reptans , we imposed selection on the proportion of flowering rosettes in the absence of competition (base population: mean = 0.391, broad-sense heritability = 0.307). We also selected on the plasticity in this trait in response to competition with a naturally coexisting grass in a parallel experiment (base population: 14% lower mean in the presence of competition, broad-sense heritability = 0.072). After two generations of bidirectional selection, the proportion of flowering rosettes was 26% higher in the high line than in the low line (realized heritability ± SE = 0.205 ± 0.017). Moreover, genotypes of the high line had 11% fewer carpels per flower, a 22% lower proportion of rooted rosettes, and a 39% smaller average distance between rosettes within a clone. In the second experiment, we found no significant responses to selection for high and low plasticity in the proportion of flowering rosettes (realized heritability ± SE =–0.002 ± 0.013). Our study indicates a high heritability and potential for further evolution of the proportion of flowering rosettes in R. reptans , but not for its plasticity, which may have been fixed by past evolution at its current level. Moreover, our results demonstrate strong genetic correlations between allocation to sexual reproduction and other clonal life-history characteristics.     

Costs of plasticity in foraging characteristics of the clonal plant Ranunculus reptans

In clonal plants, evolution of plastic foraging by increased lengths of leaves and internodes under unfavourable conditions may be constrained by costs and limits of plasticity. We studied costs and limits of plasticity in foraging characteristics in 102 genotypes of the stoloniferous herb Ranunculus reptans. We grew three replicates of each genotype with and three without competition by the naturally co-occuring grass Agrostis stolonifera. We used regression and correlation analyses to investigate potential costs of plasticity in lengths of leaves and stolon internodes, developmental instability costs of these traits, and a developmental range limit of these traits. We used randomization procedures to control for spurious correlations between parameters calculated from the same data. Under competition the number of rosettes, rooted rosettes, and flowers was 58%, 40%, and 61% lower, respectively, than in the absence of competition. Under competition lengths of leaves and stolon internodes were 14% and 6% smaller, respectively, than in the absence of competition. We detected significant costs of plasticity in stolon internode length in the presence of competition when fitness was measured in terms of the number of rosettes and the number of flowers (selection gradients against plasticity were 0.250 and 0.214, respectively). Within-environment variation (SD) in both foraging traits was not positively correlated with the corresponding plasticity, which indicates that there were no developmental instability costs. More plastic genotypes did not have less extreme trait values than less plastic genotypes for both foraging traits, which indicates that there was no developmental range limit. We conclude that in R. reptans costs of plasticity more strongly constrain evolution of foraging in the horizontal plane (i.e., stolon internode length) than in the vertical plane (i.e., leaf length).     

Effects of intraspecific competition on size variation and reproductive allocation in a clonal plant

Clonal plants grow in diameter rather than height, and therefore competition among genets is likely to be symmetric and to result in smaller variation in size of genets than in non-clonal plants. Moreover, clonal plants can reproduce both sexually and vegetatively. We studied the effects of density on the size of rosettes and of clones, variation in the size of rosettes and of clones, and allocation to sexual and vegetative reproduction in the clonal herb Ranunculus reptans . We grew plants from an artificial population of R. reptans in 32 trays at two densities. After four months, differences in density were still apparent, although clones in the low-density treatment had on average 155% more rosettes and 227% more rooted rosettes than clones in the high-density treatment. The coefficient of variation of these measures of clone size was 15% and 83% higher, respectively, in the low-density treatment. This indicates that intraspecific competition among clones of R. reptans is symmetric and increases the effective population size. Rooted rosettes were larger and varied more in size in the low-density treatment. The relative allocation of the populations to sexual and to vegetative reproduction was 19% and 13% higher, respectively, in the high-density treatment. Moreover, seeds produced in the high-density treatment had a 24% higher mass and a 7% higher germination percentage. This suggests that with increasing density, allocation to sexual reproduction increases more than allocation to vegetative reproduction in R. reptans , which corresponds to the response of some other species with a spreading growth form but not of species with a compact growth form. We conclude that intraspecific competition is an important factor in the life-history evolution of R. reptans because intraspecific competition affects its clonal life-history traits and may affect evolutionary processes such as genetic drift and selection through its effect on the effective population size.     

Effects of four generations of density-dependent selection on life history traits and their plasticity in a clonally propagated plant

Life history evolution of many clonal plants takes place with long periods of exclusively clonal reproduction and under largely varying ramet densities resulting from clonal reproduction. We asked whether life history traits of the clonal herb Ranunculus reptans respond to density-dependent selection, and whether plasticity in these traits is adaptive. After four generations of exclusively clonal propagation of 16 low and 16 high ramet-density lines, we studied life history traits and their plasticities at two test ramet-densities. Plastic responses to higher test-density consisted of a shift from sexual to vegetative reproduction, and reduced flower production, plant size, branching frequency, and lengths of leaves and internodes. Plants of high-density lines tended to have longer leaves, and under high test-density branched less frequently than those of low-density lines. Directions of these selection responses indicate that the observed plastic branching response is adaptive, whereas the plastic leaf length response is not. The reverse branching frequency pattern at low test-density, where plants of high-density lines branched more frequently than those of low-density lines, indicates evolution of plasticity in branching. Moreover, when grown under less stressful low test-density, plants of high-density lines tended to grow larger than the ones of low-density lines. We conclude that ramet density affects clonal life-history evolution and that under exclusively clonal propagation clonal life-history traits and their plasticities evolve differently at different ramet densities.     

Selection on phenotypic plasticity of morphological traits in response to flooding and competition in the clonal shore plant Ranunculus reptans

Adaptive evolution of phenotypic plasticity requires that plastic genotypes have the highest global fitness. We studied selection by spatial heterogeneity of interspecific competition and flooding, and by temporal heterogeneity of flooding on morphological plasticity of 52 genotypes of the clonal shore plant Ranunculus reptans. Competition reduced clone size, rosette size, leaf length and stolon internode thickness. Flooding had similar effects and reduced competition. Differences in selection between environments imply potential for either local adaptation or for indirect evolution of phenotypic plasticity. We also detected direct selection for plastic reductions in internode length in response to flooding and for a plastic increase in internode length in response to competition. Plastic responses of some morphological traits to flooding were in line with selection thereon, suggesting that they indeed are adaptive and might have evolved in response to direct selection on plasticity.     

On the evolution of clonal plant life histories

Clonal plant life histories are special in at least four respects: (1) Clonal plants can also reproduce vegetatively, (2) vegetative reproduction can be realised with short or long spacers, (3) and it may allow to plastically place vegetative offspring in benign patches. (4) Moreover, ramets of clonal plants may remain physically and physiologically integrated. Because of the apparent utility of such traits and because ecological patterns of distribution of clonal and non-clonal plants differ, adaptation is a tempting explanation of observed clonal life-history variation. However, adaptive evolution requires (1) heritable genetic variation and (2) a trait effect on fitness, and (3) it may be constrained if other evolutionary forces are overriding selection or by constraints, costs and trade-offs. (1) The few studies undertaken so far reported broad-sense heritability for clonal traits. Variation in selectively neutral genetic markers appears as pronounced in populations of clonal as non-clonal plants. However, neutral markers may not reflect heritable variation of life-history traits. Moreover, clonal plants may have been sampled at larger spatial scales. Empirical information on the contribution of somatic mutations to heritable variation is lacking. (2) Clonal life-history traits were found to affect fitness. However, much of this evidence stems from artificial rather than natural environments. (3) The relative importance of gene flow, inbreeding, and genetic drift, compared with selection, in the evolution of clonal life histories is hardly explored. Benefits of clonal life-history traits were frequently studied and found. However, there is also evidence for constraints, trade-offs, and costs. In conclusion, though it is very likely, that clonal life-history traits are adaptive, it is neither clear to which degree this is the case, nor which clonal life-history traits constitute adaptations to which environmental factors. Moreover, evolutionary interactions among clonal life-history traits and between clonal and non-clonal ones, such as the mating system, are not well explored. There remains much interesting work to be done in this field – which will be particularly interesting if it is done in the field.     

Three generations under low versus high neighborhood density affect the life history of a clonal plant through differential selection and genetic drift

We tested whether neighborhood density affects the clonal life history of the stoloniferous plant Ranunculus reptans through selection and genetic drift. After three generations of sexual reproduction of 16 low- and 16 high-density lines, we studied traits related to growth form and reproduction in a common competition free environment. A 7.7% lower branching frequency and slightly longer internodes indicated an evolutionary shift towards a less compact growth form under high neighborhood density, but because stolons grew also more vertically, horizontal spread per ramet was slightly decreased. Neighborhood density had no directional effects on the evolution of allocation to sexual and vegetative reproduction in R. reptans . Variation among replicated high-density lines was significantly lower than among replicated low-density lines in both growth form and reproductive characteristics, indicating less pronounced genetic drift under high neighborhood density. This study demonstrates that a clonal plant can respond to selection imposed by neighborhood density. Moreover, it shows that the effect of random genetic drift increases with decreasing neighborhood density. In a declining species, such as R. reptans in central Europe, this may lower the potential for adaptive evolutionary change and increase extinction risk.     

Genetic Allee effects on performance, plasticity and developmental stability in a clonal plant

Negative effects of small population size on fitness, so-called Allee effects, may threaten population persistence even in intact habitat remnants. We studied genotypes of 14 isolated populations of the clonal plant Ranunculus reptans, for which molecular genetic (RAPD-) variability is higher for large than for small populations. In a competition-free greenhouse environment vegetative offspring of genotypes from large populations produced more rosettes and flowers, indicating higher fitness. Within-genotype coefficients of variation in performance traits, indicating developmental instability, were lower for genotypes from populations with higher RAPD-variability. In competition with a taller grass, we found relative reduction in leaf length less pronounced for plants from large populations, suggesting higher adaptive plasticity. Our experimental study of a plant with predominantly vegetative reproduction suggests, that negative genetic effects of recent habitat fragmentation, which so far rather were expected in plants with frequent sexual reproduction, are more severe and more common than previously acknowledged.     

Genetic Basis and Selection for Life-History Trait Plasticity on Alternative Host Plants for the Cereal Aphid Sitobion avenae

Sitobion avenae (F.) can survive on various plants in the Poaceae, which may select for highly plastic genotypes. But phenotypic plasticity was often thought to be non-genetic, and of little evolutionary significance historically, and many problems related to adaptive plasticity, its genetic basis and natural selection for plasticity have not been well documented. To address these questions, clones of S. avenae were collected from three plants, and their phenotypic plasticity under alternative environments was evaluated. Our results demonstrated that nearly all tested life-history traits showed significant plastic changes for certain S. avenae clones with the total developmental time of nymphs and fecundity tending to have relatively higher plasticity for most clones. Overall, the level of plasticity for S. avenae clones’ life-history traits was unexpectedly low. The factor ‘clone’ alone explained 27.7–62.3% of the total variance for trait plasticities. The heritability of plasticity was shown to be significant in nearly all the cases. Many significant genetic correlations were found between trait plasticities with a majority of them being positive. Therefore, it is evident that life-history trait plasticity involved was genetically based. There was a high degree of variation in selection coefficients for life-history trait plasticity of different S. avenae clones. Phenotypic plasticity for barley clones, but not for oat or wheat clones, was frequently found to be under significant selection. The directional selection of alternative environments appeared to act to decrease the plasticity of S. avenae clones in most cases. G-matrix comparisons showed significant differences between S. avenae clones, as well as quite a few negative covariances (i.e., trade-offs) between trait plasticities. Genetic basis and evolutionary significance of life-history trait plasticity were discussed.     

Phenotypic plasticity and specialization in clonal versus non-clonal plants: A data synthesis

Reproductive strategies can be associated with ecological specialization and generalization. Clonal plants produce lineages adapted to the maternal habitat that can lead to specialization. However, clonal plants frequently display high phenotypic plasticity (e.g. clonal foraging for resources), factors linked to ecological generalization. Alternately, sexual reproduction can be associated with generalization via increasing genetic variation or specialization through rapid adaptive evolution. Moreover, specializing to high or low quality habitats can determine how phenotypic plasticity is expressed in plants. The specialization hypothesis predicts that specialization to good environments results in high performance trait plasticity and specialization to bad environments results in low performance trait plasticity. The interplay between reproductive strategies, phenotypic plasticity, and ecological specialization is important for understanding how plants adapt to variable environments. However, we currently have a poor understanding of these relationships. In this study, we addressed following questions: 1) Is there a relationship between phenotypic plasticity, specialization, and reproductive strategies in plants? 2) Do good habitat specialists express greater performance trait plasticity than bad habitat specialists? We searched the literature for studies examining plasticity for performance traits and functional traits in clonal and non-clonal plant species from different habitat types. We found that non-clonal (obligate sexual) plants expressed greater performance trait plasticity and functional trait plasticity than clonal plants. That is, non-clonal plants exhibited a specialist strategy where they perform well only in a limited range of habitats. Clonal plants expressed less performance loss across habitats and a more generalist strategy. In addition, specialization to good habitats did not result in greater performance trait plasticity. This result was contrary to the predictions of the specialization hypothesis. Overall, reproductive strategies are associated with ecological specialization or generalization through phenotypic plasticity. While specialization is common in plant populations, the evolution of specialization does not control the nature of phenotypic plasticity as predicted under the specialization hypothesis.     

Demographic vulnerability of the clonal and endangered meadow thistle

For effective management of endangered species it is pivotal to understand why a species is endangered and which key life cycle components are involved in its response to environmental changes. Our objective was to investigate the response of rosettes of the redlisted clonal herb Cirsium dissectum to anthropogenic nutrient enrichment, which threatens its populations, and the consequences of these responses for its population dynamics. We constructed matrix population models with demographic data from three populations and four annual transitions and we decomposed the spatiotemporal variation in projected population growth rates into contributions from life cycle components. These patterns were compared with below-ground rosette dynamics in different fields, and with the below- and above-ground rosette dynamics in a garden experiment with nutrient enrichment and competing grasses. The decomposition analysis revealed that increased clonal rosette formation and decreased rosette survival were driving the spatial variation in the population growth rate. Excavating the below-ground rhizome network revealed a higher rosette turn-over in experimentally fertilized garden plots, which not only resulted in increased plot-level extinction, but also in increased spread of the clonal offspring. This supported the observed trend among field populations: rosette formation trades off with rosette survival. Surviving seedlings were only found in areas where the topsoil had been removed. The endangered C. dissectum is vulnerable when its habitat becomes more productive, because this species does not have the necessary capability to build up biomass. Small-scale disturbances such as created by sod-cutting or trampling cattle are essential for seedling establishment and necessary to render the explorative strategy of rhizomatous clonal spread successful.     

Temperature-dependent internode elongation in vegetative plants of Arabidopsis thaliana lacking phytochrome B and cryptochrome 1

 Vegetative plants of Arabidopsis thaliana (L.) Heynh. form a compact rosette of leaves in which internode growth is virtually arrested. Rapid extension of the internodes occurs after flower buds are present in the reproductive apex. Under natural radiation, continuous light from fluorescent lamps, or short photoperiods of light from fluorescent lamps, plants of the phyB cry1 double mutant (lacking both phytochrome B and cryptochrome 1) did not form normal rosettes because all the internodes showed some degree of elongation. Internode elongation was weak in the phyB single mutant and absent in the cry1 mutant, indicating redundancy between phytochrome B and cryptochrome 1. The absence of phytochrome A caused no effects. The failure to form normal rosettes was conditional because internode elongation was arrested at low temperatures in all the mutant combinations. In contrast, the temperature dependence of phytochrome B and cryptochrome 1 effects on hypocotyl growth was weak. The elongation of the internodes in phyB cry1 was not accompanied by early flowering as showed by the lack of effects on the final number of leaves. Apex dissection indicated that in phyB cry1 double mutants internode elongation anticipated the transition from the vegetative to the reproductive stage. Thus, stem growth in Arabidopsis thaliana is not fully dependent on the program of reproductive development. Received: 2 June 1999 / Accepted: 13 August 1999     

林下和林窗内绢毛匍匐委陵菜的克隆生长和克隆形态

 为了验证绢毛匍匐委陵菜(Potentilla reptans var. sericophylla)林窗和林下种群间的行为差异是完全由表型可塑性引起,还是局部分化的结果,将生长在北京东灵山油松（Pinus tabulaeformis）林林窗和林下的绢毛匍匐委陵菜,进行生境间的交互移植-重植野外生态实验。研究结果表明,实验植物的叶片长度、叶片宽度、叶柄长度和匍匐茎节间长度等克隆形态特征在两生境间无差异。两个来源的植株,其基株生物量、基株分株数和基株匍匐茎总长度等克隆生长特征在林下生境中都比在林窗生境中小,表现出显著的可塑性。所研究的克隆形态特征和克隆生长特征及其可塑性在不同生境来源的实验植物间没有差异。绢毛匍匐委陵菜克隆形态特征和克隆生长特征及其可塑性在林下和林窗生境间没有发生局部分化,林窗为其较适生境,克隆生长特征的可塑性对绢毛匍匐委陵菜利用生境异质性可能具有重要意义     

Regeneration growth of the invasive clonal forb Rorippa austriaca (Brassicaceae) in relation to fertilization and interspecific competition

A special type of clonal growth, spread by lateral roots, ishypothesized to be a favourable trait of invasive, opportunistic plant speciesof disturbed habitats. We tested this hypothesis for the invasive forbRorippa austriaca (Brassicaceae). Regenerationfrom root fragments, subsequent vegetative spread and allocation patterns inrelation to varied nutrient supply and intensity and pattern of interspecificcompetition were analyzed in container experiments. Regeneration success fromroot fragments was 100% and clonal spread was rapid but vegetativeperformance was strongly reduced under unfertilized conditions and,particularly, when interspecific competition was present. While the ratio ofabove- to belowground bio-mass did not differ considerably betweentreatments, R. austriaca allocated a high amount ofresources to belowground growth resulting in low aboveground but highbelowground biomass at harvest time relative to the matrix vegetation.Differences in shoot number or biomass between simulated gaps and denselyvegetated quadrants in the containers were (relatively) weak.Reproductive effort was less reduced under low resource levels, and the clonesdid not set seed at all, irrespective of the treatment. Our results show thatclonal growth by lateral roots and plasticity in clonal growth patterns inR. austriaca promote both exploitation of gaps andnutrient-rich microsites and resistance to competitors. Such plasticity,combined with its ability to regenerate from widely-dispersed rootfragments, contribute to the ability of the species to invade and persistwithindisturbed and spatially heterogeneous habitats.     

Estimation of heritability using variety trials data from incomplete blocks

An assessment of the heritability of a trait is useful in formulating a breeding strategy for crop improvement. We have considered the estimation of broad-sense heritability from a single-location trial and from multi-locational trials conducted in incomplete blocks. Using residual maximum likelihood estimates of variance components, we estimated the heritability and obtained expressions for the estimate of its bias and its standard error. The estimation procedure is illustrated for 25 barley genotypes evaluated at four locations in West Asia and North Africa during 1992.     

Variation in petiole and internode length affects plant performance in <Emphasis Type="Italic">Trifolium repens</Emphasis> under opposing selection regimes

We studied the effects of genotypic and plastic variation in vertical and horizontal spacer lengths on plant performance in a stoloniferous herb subjected to opposing selection regimes. We hypothesized that longer vertical structures are beneficial if plants are subjected to competition, but they should negatively affect plant performance if plants are exposed to aboveground disturbance. To test these hypotheses we subjected 34 genotypes of Trifolium repens to competition and disturbance treatments. Competition was imposed by a grass canopy consisting of Lolium perenne, and disturbance was simulated by regularly clipping the target plants and all the surrounding vegetation at 1 cm above soil level. Conform to our hypothesis, genotypes with longer vertical structures (petioles) produced fewer ramets than genotypes with shorter petioles in the disturbance treatment. However, genotypes with longer petioles did not perform better under competition than genotypes with shorter petioles. Genotypes with highly plastic vertical structures tended to produce more shoot mass under competition, and they produced fewer ramets if subjected to disturbance. Unexpectedly, horizontal structures (stolon internodes) expanded in response to competition which, furthermore, was associated with enhanced plant performance. However, producing longer internodes is inherently associated with costs in terms of increased resource allocation to the longer structures, but not to benefits in terms of increased resource capture. Positive correlations among the length and plasticity of vertical and horizontal structures may explain the apparent positive effect of producing longer internodes on plant performance. Our data thus support the notion that trait correlations may weaken selective forces acting on a focal trait in a specific environment if opposing selection pressures act on genetically correlated traits.     

The effects of reproductive specialization on energy costs and fitness genetic variances in cyclical and obligate parthenogenetic aphids

Organisms with coexisting sexual and asexual populations are ideal models for studying the consequences of either reproductive mode on the quantitative genetic architecture of life-history traits. In the aphid Rhopalosiphum padi, lineages differing in their sex investment coexist but all share a common parthenogenetic phase. Here, we studied multiple genotypes of R. padi specialized either for sexual and asexual reproduction and compared their genetic variation in fitness during the parthenogenetic phase. Specifically, we estimated maintenance costs as standard metabolic rate (SMR), together with fitness (measured as the intrinsic rate of increase and the net reproductive rate). We found that genetic variation (in terms of broad-sense heritability) in fitness was higher in asexual genotypes compared with sexual genotypes. Also, we found that asexual genotypes exhibited several positive genetic correlations indicating that body mass, whole-animal SMR, and apterous individuals production are contributing to fitness. Hence, it appears that in asexual genotypes, energy is fully allocated to maximize the production of parthenogenetic individuals, the simplest possible form of aphid repertoire of life-histories strategies.     

Fire adaptation in Neblinaria celiae (Theaceae), a high-elevation rosette shrub endemic to a wet equatorial tepui

Summary Neblinaria celiae (Theaceae), a rosette shrub endemic to the exceedingly rainy summit of remote Cerro de la Neblina in southern Venezuela, has a previously undescribed set of adaptations to fire. Its growth form entails sparse branching, massive terminal leaf rosettes, and thick bark. It is highly fire-tolerant, with a survival rate of 93% in a stand recently ignited by lightning, vs. 0% in seven co-occurring woody species. Survival increases sharply with rosette height, favoring a sparsely branched habit that would maximize the rate of upward growth through the sparse fuel layer supported by a sterile substrate. Thick bark and massive rosettes help protect cambial and foliar meristems from brief exposure to high temperatures. Rosettes on shorter plants are exposed to greater damage from fire near the ground and, as expected, are bigger and impound more rainwater; the greater number of leaves nearly balances the greater leaf mortality caused by fire. We relate Neblinaria's growth form to its dominance atop Neblina, to a general model for the evolution of sparse branching, and to the evolution of growth form in other tepui plants.     

光照强度和基质养分对匍匐茎草本金戴戴克隆生长和克隆形态的影响

在深度遮光(光照强度为高光条件的6.25%,约为自然光照的5.3%)或低养分条件下,金戴戴(Halerpestes ruthenica Ovcz.)生物量、初级分株叶面积、分株总数、匍匐茎总数和总长度均显著减小,而比节间长和比叶柄长显著增加.在低养分条件下,金戴戴匍匐茎平均节间长显著增加,而匍匐茎分枝强度和分株数显著减小.这些结果与克隆植物觅食模型相符合,表明当生长于异质性生境中,金戴戴可能通过以克隆生长和克隆形态的可塑性实现的觅养行为来增加对养分资源的摄取.在深度遮光条件下,金戴戴平均间隔子长度(即平均节间长和平均叶柄长)均显著减小.这一结果与以往实验中匍匐茎草本间隔子对中度和轻度遮光(光照强度为高光条件的13%～75%,>10%的自然光照)的反应不同.这表明,在深度遮光条件下匍匐茎克隆植物可能不发生通过间隔子可塑性实现的觅光行为.光照强度和基质养分条件的交互作用对许多性状如总生物量、匍匐茎总数和总长度、二级和三级分株数、分株总数、初级分株叶面积以及分枝强度均有十分显著的效应.在高光条件下,基质养分对这些性状有十分显著的影响;而在低光条件下,基质养分条件对这些性状不产生影响或影响较小.这表明,光照强度影响金戴戴对基质养分的可塑性反应.在深度遮光或低养分条件下,金戴戴可能通过减小匍匐茎节间粗度(增加比节间长)来增加或维持其相对长度,从而更有机会逃离资源丰度低的斑块.     

The movement ecology of seagrasses

A movement ecology framework is applied to enhance our understanding of the causes, mechanisms and consequences of movement in seagrasses: marine, clonal, flowering plants. Four life-history stages of seagrasses can move: pollen, sexual propagules, vegetative fragments and the spread of individuals through clonal growth. Movement occurs on the water surface, in the water column, on or in the sediment, via animal vectors and through spreading clones. A capacity for long-distance dispersal and demographic connectivity over multiple timeframes is the novel feature of the movement ecology of seagrasses with significant evolutionary and ecological consequences. The space–time movement footprint of different life-history stages varies. For example, the distance moved by reproductive propagules and vegetative expansion via clonal growth is similar, but the timescales range exponentially, from hours to months or centuries to millennia, respectively. Consequently, environmental factors and key traits that interact to influence movement also operate on vastly different spatial and temporal scales. Six key future research areas have been identified.