Importance of starting points in heterogeneous environments: interactions between two clonal plants with contrasting spatial architectures

Aims Plants can benefit from heterogeneous environments via disproportionately increasing resource harvesting in resource-rich patches.Their initial growing positions with respect to resource patches may thus have important influences on their performance and relative competitive ability.Such impacts may differ between species with contrasting spatial architectures.However,the potential influence of initial growing positions in heterogeneous environment on plant growth and competition has largely been ignored.Methods We grew the phalanx plant Carex neurocarpa and the guerrilla plant Bolboschoenus planiculmis alone or in competition in a heterogeneous environment consisting of high-and low-nutrient soil patches.In treatments without competition,one ramet of each species was grown in either a high-or a low-nutrient patch in the heterogeneous environment.In treatments with competition,a ramet of the target species was grown in either a high-or a low-nutrient patch,and a ramet of the competitor species was grown in the same patch as the target species or an adjacent patch with a different nutrient level.Important Findings Without competition C.neurocarpa produced more biomass and ramets when initially grown in a high-nutrient patch than when initially grown in a low-nutrient patch.With competition,these differences disappeared.Consequently,competitive intensity on C.neurocarpa was higher when it initially grew in a high-nutrient patch than when it initially grew in a low-nutrient patch.These impacts were independent of the initial position of its competitor.By contrast,the initial positions of B.planiculmis did not influence its growth or competitive response.Therefore,in heterogeneous environments,initial growing positions of clonal plants may influence their performance in competition-free environments and may also affect their relative competitive ability,and these effects may depend on spatial architecture of the plants.

Role of Clonal Integration among Different Environmental Conditions (A Review)

Most of the ecosystems are dominated by clonal species. The most unique feature of clonal plants is their capability for clonal integration (translocation of vital resources among connected ramets), implying that integration may play an essential role in their success. However, a general effect of clonal integration on plant performance is lacking. We conducted a text review on the effects of clonal integration on different habitats and species. Overall, clonal integration increased performance of clonal plants in different habitats. However clonal integration has also some limitations under stressful environments. Benefits of clonal integration may lack somehow when environmental stress increases. But connected ramets placed in unfavorable patches benefited more from integration compared to severed ramet placed in nutrient rich patches. Climate change and temperature increase have positive effects on biomass of clonal species.

Study on Clonal Growth Pattern of Sabina vulgaris in Mu Us Sandland

The clonal growth pattern of Sabina vulgaris, a coniferous clonal plant in Mu Us sandland, Inner Mongolia Autonomous Region was surveyed. The results showed that with the stolon extending, internode length and branching angle decreased, the branching intensity increased gradually within the 3 m range from the edge of the shrub to its center along the stolon. Internode length, branching intensity and branching angle were 5.9 cm, 4.4 and 55.3°in the shrub, and 1.6 cm, 13.7 and 38.3°at the edge of the shrub, respectively. The clonal architecture exhibited plasticity. The internode length, branching intensity and ramet length changed with an exponential model with extention of the stolon. The stolon of S. vulgaris was monopodial branching, and each ramet should possess more than 3 adventitious roots. Ramets could take on the phenomenon of 搒elf-thinning?with clonal growth. There was a prior grade in allocation of the nutrients gained from heterogeneous space. The clonal architecture of S. vulgaris was the 搈ixed?type.

Clonal Transgenerational Effects Transmit for Multiple Generations in a Floating Plant

Environmental conditions of a parent plant can influence the performance of their clonal offspring,and such clonal transgenerational effects may help offspring adapt to different environments.However,it is still unclear how many vegetative generations clonal transgenerational effects can transmit for and whether it depends on the environmental conditions of the offspring.We grew the ancestor ramets of the floating clonal plant Spirodela polyrhiza under a high and a low nutrient level and obtained the so-called 1^(st)-generation offspring ramets of two types(from these two environments).Then we grew the 1^(st)-generation offspring ramets of each type under the high and the low nutrient level and obtained the so-called 2^(nd)-generation offspring ramets of four types.We repeated this procedure for another five times and analyzed clonal transgenerational effects on growth,morphology and biomass allocation of the 1^(st)-to the 6^(th)-generation offspring ramets.We found positive,negative or neutral(no)transgenerational effects of the ancestor nutrient condition on the offspring of S.polyrhiza,depending on the number of vegetative generations,the nutrient condition of the offspring environment and the traits considered.We observed significant clonal transgenerational effects on the 6^(th)-generation offspring;such effects occurred for all three types of traits(growth,morphology and allocation),but varied depending on the nutrient condition of the offspring environment and the traits considered.Our results suggest that clonal transgenerational effects can transmit for multiple vegetative generations and such impacts can vary depending on the environmental conditions of offspring.

Resource Sharing by the Different Clonal White Clover Plant(Trifolium repens L.)

Resource sharing in the different parts of the clonal white clover plant (Trifolium repens L.) was investigated in this study. The results showed that the integration of water and nitrogen metabolism occured among parental, young and old branches of a plant when water passed through its stolons. Under the sufficient supply of light, water and nutrients from soil the newly developed branches showed similar capacity in taking in sufficient water and nitrogen to that of old branches. The development of the new branches did not affect the growth of the old and parental branches attached. The results also indicated that energy values in roots was significantly related with plant age, however, the energy values were various in the different parts of a plant.

Phenotypic Plasticity in Response to the Heterogeneous Water Supplyin the Rhizomatous Grass Species, Calamagrostis epigejosin the Mu Us Sandy Land of China

Calamagrostis epigejos (L.) Roth. is a perennial grass with slender and long rhizome segments between interconnected neighbor ramets. To investigate the phenotypic plasticity in response to the heterogeneous soil water supply, ramet pairs of the species were subjected to heterogeneous water supply by which either mother ramets or daughter ramets were in high or low soil water supply, respectively, in the Maowusu (Mu Us) Sandy Land of Nei Mongol. The results showed that the phenotypic characteristics of the individual ramets of C epigejos were greatly influenced by the heterogeneous water supply. The ramets treated with high water supply significantly produced more new rhizomes and more offspring (ramets), and accumulated more shoot biomass, and allocated more biomass to their shoots than those treated with low water supply. In comparison with the daughter ramets in homogeneous soil water supply, phenotypic characteristics, in terms of new rhizome growth, the production of new offspring, and the biomass allocation pattern, of the daughter ramets within the pairs of the species were not significantly changed, no matter that high or low soil water supply to mother ramets. The phenotypic responses of mother ramets to soil water supply were similar to those of daughter ramets. From these results, it is inferred that the interconnected ramets of C epigejos response phenotypically to their local soil water rather than to the soil water experienced by the interconnected ramets. The interconnected ramets of C epigejos might be independent of each other in water relationship, although they are physically interconnected with rhizome segments. The physiological independence of interconnected ramets might facilitate the risk spreading and thus enhance the genet survivorship under the frequent drought stresses in Mu Us Sandland.

Architectural Plasticity in Response to Soil Moisture in the Stoloniferous Herb, Duchesnea indica

Architectural plasticity of clonal plants may enhance exploitation of soil moisture heterogeneity by the plants. The plasticity of clonal architecture in response to soil moisture in the stoloniferous herb, Duchesnea indica Focke, was investigated in an experiment with different soil moisture contents as treatments, i.e. 40%, 60%, 80%, 100% of the maximum moisture content of soil (MMCS). As soil moisture content increased, the spacer length, ramet density, branching intensity and branching angle of D. indica plants changed by quadratic curve. And the optimum habitat for the plants was at 80% of the MMCS. This architectural plasticity in D. indica was simulated through the Dynamic Logistic Model. The imitative effect was statistically satisfactory. Its architectural plasticity observed here may allow the species to show foraging behavior in its habitat where soil moisture is patchily distributed.

Simulating the evolution of a clonal trait in plants with sexual and vegetative reproduction

Aims Phenotypic optimality models neglect genetics.However,especially when heterozygous genotypes are fittest,evolving allele,genotype and phenotype frequencies may not correspond to predicted optima.This was not previously addressed for organisms with complex life histories.Methods Therefore,we modelled the evolution of a fitness-relevant trait of clonal plants,stolon internode length.We explored the likely case of an asymmetric unimodal fitness profile with three model types.In constant selection models(CSMs),which are gametic,but not spatially explicit,evolving allele frequencies in the one-locus and fiveloci cases did not correspond to optimum stolon internode length predicted by the spatially explicit,but not gametic,phenotypic model.This deviation was due to the asymmetry of the fitness profile.Gametic,spatially explicit individual-based(SEIB)modeling allowed us relaxing the CSM assumptions of constant selection with exclusively sexual reproduction.Important findings For entirely vegetative or sexual reproduction,predictions of the gametic SEIB model were close to the ones of spatially explicit nongametic phenotypic models,but for mixed modes of reproduction they approximated those of gametic,not spatially explicit CSMs.Thus,in contrast to gametic SEIB models,phenotypic models and,especially for few loci,also CSMs can be very misleading.We conclude that the evolution of traits governed by few quantitative trait loci appears hardly predictable by simple models,that genetic algorithms aiming at technical optimization may actually miss the optimum and that selection may lead to loci with smaller effects in derived compared with ancestral lines.

Differential Effects of Ammonium and Nitrate on Growth Performance of Glechoma longituba under Heterogeneous Cd Stress

Water,minerals,nutrients,etc.,can be shared by physiological integration among inter-connected ramets of clonal plants.Nitrogen plays an important role in alleviating cadmium(Cd)stress for clonal plants.But how different forms of nitrogen affect growth performance of clonal plants subjected to heterogeneous Cd stress still remains poorly understood.A pot experiment was conducted to investigate the differential effects of ammonium and nitrate on growth performance of Glechoma longituba under heterogeneous Cd stress.In the experiment,parent ramets of Glechoma longituba clonal fragments were respectively supplied with modified Hoagland solution containing 7.5 mM ammonium,7.5 mM nitrate or the same volume of nutrient solution without nitrogen.Cd solution with different concentrations(0,0.1 or 2.0 mM)was applied to offspring ramets of the clonal fragments.Compared with control(N-free),nitrogen addition to parent ramets,especially ammonium,significantly improved antioxidant capacity[glutathione(GSH),proline(Pro),peroxidase(POD,)superoxide dismutase(SOD)and catalase(CAT)],PSII activity[maximum quantum yield of PSII(F_(v)/F_(m))and effective quantum yield of PSII(ΦPSII)],chlorophyll content and biomass accumulation of the offspring ramets suffering from Cd stress.In addition,negative effects of nitrate on growth performance of whole clonal fragments were observed under Cd stress with high concentration(2.0 mM).Transportation or sharing of nitrogen,especially ammonium,can improve growth performance of clonal plants under heterogeneous Cd stress.The experiment provides insight into transmission mechanism of nitrogen among ramets of clonal plants suffering from heterogeneous nutrient supply.Physiological integration might be an important ecological strategy for clonal plants adapting to heterogeneous environment stress conditions.

Quantitative Estimates of Outcrossing Rates in a Natural Population of Caldesia grandis (Alismataceae)

Outcrossing rate in a natural population of Caldesia grandis was estimated by the dominant random amplified polymorphism DNA （RAPD） marker using 10 open-pollinated progeny arrays of 24 individuals. The multilocus outcrossing rate estimated based on all 25 RAPD loci was 0.872 ±0.033 and the single-locus outcrossing rate was 0.795 ±0.032. Multilocus esti- mates did not differ significantly from the single-locus estimates. The fixation index, F, in the progeny estimated from RAPD data was -0.142 ±0.000. The estimates of multilocus outcrossing rates （tm） and single-locus outcrossing rates （ts） obtained from MLDT clearly indicate that outcrossing is predominant in the open-pollinated C. grandis population. An empirical analysis suggests that 15 should be the minimum number of dominant marker loci necessary to achieve robust estimates of tm.

Soil Moisture Rather than Soil Nutrient Regulates the Belowground Bud Bank of Rhizomatous Species Psammochloa villosa in Arid Sand Dunes

In arid and semi-arid sand dune ecosystems,belowground bud bank plays an important role in population regeneration and vegetation restoration.However,the responses of belowground bud bank size and composition to sand burial and its induced changes in soil environmental factors have been rarely studied.In arid sand dunes of Northwestern China,we investigated belowground bud bank size and composition of the typical rhizomatous psammophyte Psammochloa villosa as well as three key soil environmental factors(soil moisture,total carbon and total nitrogen)under different depths of sand burial.Total buds and rhizome buds increased significantly with increasing burial depth,whereas tiller buds first increased and then decreased,with a peak value at the depth of 20-30 cm.Soil moisture increased significantly with sand burial depth,and was positively correlated with the number of all buds and rhizome buds.Soil total carbon concentration first increased and then decreased with sand burial depth,and total nitrogen concentration was significantly lower under deep sand burial than those at shallow depths,and only the number of tiller buds was positively correlated with soil total nitrogen concentration.These results indicate that soil moisture rather than soil nutrient might regulate the belowground bud bank of P.villosa,and that clonal psammophytes could regulate their belowground bud bank in response to sand burial and the most important environmental stress(i.e.,soil moisture).These responses,as the key adaptive strategy,may ensure clonal plant population regeneration and vegetation restoration in arid sand dunes.

Bioreactor technology for clonal propagation of plants and metabolite production

Plant cell culture in bioreactors is an enabling tool for large scale production of clonal elite plants in agriculture, horticulture, forestry, pharmaceutical sectors, and for biofuel production. Advantages of bioreactors for plant cell culture have resulted in various types of bioreactors differing in design, operating technologies, instrumentations, and construction of culture vessels. In this review, different types of bioreactors for clonal propagation of plants and secondary metabolites production are discussed. Mechanical and biochemical parameters associated with bioreactor design, such as aeration, flow rate, mixing, dissolved oxygen, composition of built-up gas in the headspace, and pH of the medium, are pivotal for cell morphology, growth, and development of cells within tissues, embryos, and organs. The differences in such parameters for different bioreactor designs are described here, and correlated to the plant materials that have been successfully cultured in different types of bioreactors.

Emergent plant presence and richness alter competitive interactions between two foating plants

Interactions between two plant species can be infuenced by the presence of other plant species and such an effect may change as the diversity of the other species increases.To test these hypotheses,we frst constructed aquatic communities consisting of 1,2 and 4 emergent plant species and then grew ramets of Lemna minor only,ramets of Spirodela polyrhiza only or ramets of both L.minor and S.polyrhiza within these aquatic communities.We also included controls with ramets of L.minor,S.polyrhiza or both but without any emergent plants.Biomass and number of ramets of L.minor and S.polyrhiza were signifcantly smaller with than without the emergent plants,but they did not differ among the three richness levels.The presence of S.polyrhiza did not signifcantly affect the growth of L.minor,and such an effect was not dependent on the richness of the emergent plant species.Without the emergent plant species,the presence of L.minor markedly reduced biomass(-92%)and number of ramets(-88%)of S.polyrhiza.However,such a competitive effect of L.minor on S.polyrhiza became much weaker in the presence of one emergent plant species(-46%biomass and-39%number of ramets)and completely disappeared in the presence of two or four emergent plant species.Therefore,both the presence and richness of emergent plant species can alter competitive interactions between the two duckweed species.These fndings highlight the importance of species diversity in regulating plant-plant interactions.

Adaptive plasticity in response to light and nutrient availability in the clonal plant Duchesnea indica

Phenotypic plasticity enables plants to buffer against environmental stresses and match their phenotypes to local conditions.However,consistent conclusive evidence for adaptive plasticity has only been obtained for a few traits.More studies on a wider variety of plant functional traits and environmental factors are still needed to further understand the adaptive significance of plasticity.We grew 21 genotypes of the stoloniferous clonal plant Duchesnea indica under different light and nutrient conditions,and used selection gradient analyses to test the adaptive value(benefits)of morphological and physiological plasticity responding to variation in light and nutrient availability.Plants grown in shade exhibited lower values for fitness measures(fruit number,ramet number and biomass),shortened thinner internode length and decreased adult leaf chlorophyll content,but higher petiole length,specific leaf area and old leaf chlorophyll content,than plants grown without shade.Plants grown in the low nutrient condition had shorter petiole length,thicker and smaller leaf area,lower chlorophyll content,but higher fruit number and root:shoot ratio than plants grown under the high nutrient condition.Selection gradient analyses revealed that plasticity of petiole length and old leaf chlorophyll content in response to light variation was adaptive,and plasticity of old and adult leaf chlorophyll content in response to nutrient variation was adaptive.Therefore,the adaptive value of plasticity in different traits depends on the specific ecological context.Our findings contribute to understanding the adaptive significance of phenotypic plasticity of clonal plants in response to environmental variation.

Clonal Plasticity in Response to Reciprocal Patchiness of Light and Nutrients in the Stoloniferous Herb Glechoma Iongituba L.

Interconnected ramets of clonal plants can functionally specialize in the uptake of resources from aboveground and/or underground sources. Ramet pairs of the clonal stoloniferous herb Glechoma Iongltuba L. were grown In spatially heterogeneous environments with complementary availability of light and nutrients. Plasticity with respect to root-shoot ratio, fitness-related traits （biomass, number of ramets and dry weight per ramet）, morphological traits （lamina area, root length） were experimentally examined. The aim was to understand the adaptation of G. Iongltuba to an environment with reciprocal patchiness of light and soil nutrients by plasticity In Its root-shoot ratio and clonal morphology. The results showed that the performance of ramets growing In patches with high light Intensity and low soil nutrients into the adjacent opposite patches was Increased in terms of fitness-related traits. However, the performance of those from patches with low light Intensity and high soil nutrients into the adjacent opposite patches was not changed. The root-shoot ratio and clonal morphology were plastic. Morphological traits such as lamina area and root length were altered In a way that enhanced the capture of light resources and soil nutrients. Apparent reciprocal resource transport between the ramets In an environment of reciprocal patchiness of resources can enhance the growth of ramets with complementary resource deficiencies.

Molecular Genetic Variation in a Clonal Plant Population of Leymus chinensis （Trin.） Tzvel.

Randomly amplified polymorphic DNA （RAPD） analysis was used to investigate the genetic variation among populations, between populations, and within populations, relationships between genetic distance and geographic distance, and the molecular variation and population size. The effects of geographic and genetic distances, as well as of genetic differentiation and population size, on genetic variations of Leymus chinensis （Trin.） Tzvel. are discussed. The present study showed that there was significant RAPD variation between the Baicheng region population and the Daqing region population, with a molecular variance of 6.35% （P 〈 0.04）, and for differentiation among area populations of the Daqing region, with a molecular variance of 8.78% （P 〈 0.002）. A 21.06% RAPD variation among all 16 populations among two regions was found （P 〈 0.001）, as well as 72.59% variation within populations （P 〈 0.001）. Molecular variation within populations was significantly different among 16 populations.

Effects of salinity and clonal integration on the amphibious plant Paspalum paspaloides:growth,photosynthesis and tissue ion regulation

Aims Clonal integration can increase performance of clonal plants suffer-ing from environmental stress,and clonal plants in many wetlands commonly face stress of flooding accompanied by salinity.However,few studies have tested roles of clonal integration in amphibious plants expanding from terrestrial to aquatic saline habitats.Methods Basal(older)ramets of clonal fragments of Paspalum paspaloides were grown in soil to simulate terrestrial habitats,whereas their apical(younger)ramets were placed at the surface of saline water containing 0,50,150 and 250 mmol l^(−1)NaCl to mimic different salinity levels in aquatic habitats.Stolons connecting the apical and basal ramets were either intact(connected)to allow clonal integra-tion or severed(disconnected)to prevent integration.Important Findings Increasing salinity level significantly decreased the growth of the apical ramets of P.paspaloides,and such effects on the leaf growth were much higher without than with stolon connection after 60-day treatment.Meanwhile,leaf and total mass ratios of the connected to the disconnected apical ramets were higher at high than at low saline treatments.Correspondingly,Fv/Fm and F/Fm′of the apical ramets were higher with than without stolon connection in highly saline treatments.The results suggest that clonal integration can benefit the spread of apical ramets from terrestrial soil into saline water,and that the positive effects increase with increasing salinity.However,clonal integration did not significantly affect the growth of the whole frag-ments.Due to clonal integration,Na^(+)could be translocated from the apical to the basal ramets to alleviate ion toxicity in apical ramets.Our results suggest that clonal integration benefits the expansion of P.paspaloides from terrestrial to aquatic saline habitats via maintained photosynthetic capacities and changed biomass allocation pattern.

Opposing effects of plant growth regulators via clonal integration on apical and basal performance in alligator weed

Invasive plants are a major threat to biodiversity and may adversely affect food security.Clonal integration enables the sharing of resources between connected ramets and can enhance plant performance in many invasive species.However,few studies have examined the role of clonal integration when weeds are exposed to plant growth regulators(PGRs).PGRs are used extensively in agriculture and may affect nearby weeds through soil leaching,erosion and runoff.Our aim was to investigate the effects of clonal integration on growth in a noxious weed,Alternanthera philoxeroides(alligator weed),in response to two PGRs frequently used in agriculture,gibberellins(GAs)and paclobutrazol(PAC).Ramets of A.philoxeroides were propagated in the greenhouse,and treated with PGRs.PGRs were applied to the older ramets(i.e.‘basal’part),with half of the plants having the stems between the apical(younger)and basal parts left connected,while the remaining plants had the stems between the two parts severed.Following the growing period,plants were measured for growth traits.We found that GA and PAC had contrasting effects on plant growth.GA significantly promoted above-ground growth of the apical ramets via clonal integration.Alternatively,PAC inhibited above-ground growth in the basal and apical parts,and enhanced below-ground growth of the basal and apical ramets through clonal integration.Our results highlight how clonal integration can promote growth in A.philoxeroides following the application of PGRs,which is likely an important mechanism for this species to invade new environments.

Genotypic differences in response to different patterns of clonal fragmentation in the aquatic macrophyte Pistia stratiotes

Connected individuals(ramets)of clonal plants are frequently fragmented due to disturbance,and such clonal fragmentation may influence their growth.However,it is unclear whether different patterns of clonal fragmentation produce differential effects on plant growth and whether such differences vary with genotypes.We collected one group of connected ramets of the stoloniferous floating invasive macrophyte Pistia stratiotes from each of six provinces(Guangdong,Guangxi,Hubei,Jiangsu,Sichuan and Yunnan Provinces)of China,and assumed that these ramets belonged to different genotypes.After several generations of vegetative propagation to remove potential parental effects,new ramets(hereafter referred to as parent ramets)from each of these assumed genotypes were subjected to four patterns of clonal fragmentation,i.e.,all offspring ramets produced during the experiment remained connected to their parent ramet(control),or the primary(1°),secondary(2°)or tertiary(3°)offspring ramets were disconnected from their parent ramet.We found significant growth differences among the assumed genotypes,confirming that the ramets sampled from these six provinces were from different genotypes.Compared with the control,disconnecting the 1°,2°or 3°ramets did not significantly affect biomass of the whole clone(i.e.,the parent ramet plus all offspring ramets)of P.stratiotes.Disconnecting the 1°ramets significantly increased biomass of the parental ramet of all six genotypes compared with the control.However,disconnecting the 2°or 3°ramets had little effect on biomass of the parent ramet for all genotypes except the Guangxi genotype,for which disconnecting the 2°ramets decreased biomass of the parent ramet.Disconnecting the 1°,2°or 3°ramets had no impact on biomass or number of the 1°or the 3°ramets for any of the genotypes.However,disconnecting the 1°ramets decreased biomass of the 2°ramets for the Guangdong and Yunnan genotypes,but had no effect for the other four genotypes.We conclude that different patterns of clonal fragmentation can have different impacts on parent and offspring ramets of P.stratiotes and such an impact may vary with genotypes.

Kin recognition benefits clonal offspring performance of historically grazed Leymus chinensis:insights for grassland management

Background:Individual plants can identify their neighbors and adjust their biomass investment to avoid competing with their relatives or jointly cope with external stresses.Maternal effects can improve their offspring adaptability under external stresses.However,how grazing-induced maternal effects influence plant kin interactions remain unknown.Methods:Clonal offspring of Leymus chinensis grown under multi-year grazing and non-grazing conditions were used for this study.A greenhouse experiment was conducted to evaluate the performance of focal plants in the presence of kin and stranger neighbors,with the aim of analyzing the interaction between maternal effect and kin relatedness.Results:Kin relatedness of neighboring plants affected the biomass production and allocation of focal plants,demonstrating the presence of kin recognition in L.chinensis.Moreover,grazing-induced maternal effects significantly enhanced kin recognition in the species.Consequently,the presence of stranger neighbors significantly improved the growth potential of grazed offspring.Specifically,the total biomass of clonal offspring increased by 73.1%compared to the kin group,potentially buffering grazing-induced plant productivity declines.Conclusions:This study shows that historical grazing enhances kin recognition in L.chinensis.Thus,introducing multi-genotypic plants can increase the productivity of grasslands.The findings of this study enhance our understanding of intraspecific plant–plant interactions in clonal species and provide new insights into sustainable grassland management.