Distribution of clonal growth traits among wetland habitats

Clonality resulting from the growth of specialized organs is common among plants in wetland habitats. We hypothesize that different wetland habitats select for different attributes of clonal traits. This hypothesis is based on studies of individual species but has not been previously tested at the level of habitat. We compared the functional diversity of clonal growth traits of plants in bogs, fens, wet heathlands, floodplains, river beds, open fresh water habitats, salt marshes, and open marine habitats. Clonal traits (including number of offspring, lateral spread, persistence of connections between ramets, and shoot life span) were analysed with multivariate techniques using species frequency data and with permutation tests using presence/absence data. Based on species frequencies, clonal plants in aquatic habitats (open fresh water habitats, open marine habitats, and river beds) were characterized by the abundant production of freely dispersible propagules, annual shoots, and splitting clones. Species of daily flooded salt marshes were characterized by bi-annual connections between ramets and medium dispersability. In contrast, plants in permanently wet bogs were characterized by polycyclic shoots and low offspring production. The specificity of river beds and open freshwater habitats was also confirmed by permutation tests, which gave equal weight to rare and abundant species. However, species in all other wetland habitats were characterized by the entire range of clonal traits, suggesting weak environmental filtering of analyzed traits by habitat at the present scale.     

Patterns of functional clonal traits and clonal growth modes in contrasting grasslands in the central Apennines,Italy

Abstract. Aim: Patterns of plant functional traits related to clonality (clonal growth modes; CGM) in plant communities were studied and hypotheses on the importance of the selected traits in plant communities supported by soils differing in moisture and nutrient status were tested. Material and Methods: Selected plant functional traits, such as the position of the mother‐daughter plants connections, length of spacers, frequency of multiplication, persistency of ramets connections, presence of storage organs and bud protection were studied in two contrasting plant communities (xeric and mesic abandoned pastures) typical of central Apennines, Italy. Results and Discussion: Clonality was shown to be of great importance in both mesic and xeric grasslands. The major differences between the two communities were due to the dominant CGMs: turf graminoids (having effective protection of growth meristems in dense tussocks) dominated xeric grasslands, while rhizomatous graminoids (typical of competitive resource‐rich environments) dominated mesic grasslands. Below‐ground CGOs (clonal growth organs), shorter spacers, higher multiplication potential, permanent ramet connection, large bud bank and increased importance of bud protection were found to be of importance in water stressed xeric grassland. Contrary to our expectations, the mesic (less stressed) grasslands have the higher number of clonal plants possessing storage organs.     

Clonal Growth Strategies Along Flooding and Grazing Gradients in Atlantic Coastal Meadows

Specific composition and species clonal traits were characterized along combined flooding and grazing gradients to answer two questions. i) To what extent does the interaction of flooding and grazing influence the clonal characteristics of the vegetation? ii) Are the effects of both environmental factors independent or interactive? This study was carried out in a wet meadow along the Atlantic coast (France). Three plant communities (hygrophilous, meso-hygrophilous and mesophilous) were distinguished along a flooding gradient and five levels of grazing pressure were controlled through an experimental design (from no grazing to heavy grazing). We monitored species composition and retrieved, for each species, the type of clonal growth organs (CGOs) and clonal traits from the CLO-PLA3 database. We identified two syndromes of clonal traits: ??above-ground splitters?? and ??below-ground integrators??. Clonal traits played a key role in plant assembly in the studied meadows. The interaction of both environmental factors selected for particular syndromes of clonal traits; however, flooding had a stronger filtering effect than grazing. The hygrophilous community was dominated by above-ground splitters, whereas the meso-hygrophilous vegetation was dominated by below-ground integrators. In the mesophilous community, clonal composition was the most diverse and shared clonal traits with the vegetation of both the hygrophilous and meso-hygrophilous communities. Grazing impact on CGOs and clonal traits differed between plant communities, i.e., the effect of grazing was modulated by the flooding regime. This study confirmed that vegetation responses to grazing might depend on the pool of traits, primarily filtered by environmental factors such as flooding.     

Classifying clonal growth forms based on vegetative mobility and ramet longevity: a whole community analysis

We measured rhizome branching, clonal mobility, and ramet longevity of 98 meadow plant species. A cluster analysis applied to this dataset revealed nine clonal growth types that differ mainly by the ramet lifespan and vegetative mobility. Then we compared the abundance of these groups of clonal species between the three following plant communities: (1) open, (2) restored and (3) overgrown wooded meadows in the Laelatu-Nehatu-Puhtu Nature Reserve, Estonia. This is the first study where the quantitative values of belowground clonal traits have been measured for all species of a species-rich community. We show that species with annual ramets and with a low vegetative mobility were most abundant in open grasslands. The relative abundance of perennial species with annual ramets was positively correlated with shoot density and species diversity, indicating that high ramet turnover rates combined with a high genet longevity can positively affect species coexistence in meadow communities. Hence, this study provides evidence for the fact that the average values of clonal life-history parameters differ between these communities. Herb communities under forest canopy consist, in average, of species with ramets that live longer and are clonally more mobile than in the communities of open sites.     

A Comparative Study on Photosynthesis and Water Use Efficiency Between Clonal and Non-clonal Plant Species Along the Northeast China Transect (NECT)

Net photosynthesis (P n), transpiration (E), stomatal conductance (g s) and water use efficiency (WUE) of more than 218 species belonging to two different reproductive functional types, i.e. clonal (115 species) and non-clonal species (103 species), along the 1 670 km Northeast China Transect (NECT) were analyzed. The results showed that P n and WUE appeared to be lower in the east and west ends of NECT, with peaks in the middle. Transpiration was found to be higher in the west end, where most temperate desert species were distributed. On the same site, most clonal species showed higher P n and related physiological variables than non-clonal species. For different growth forms over NECT, e.g. forest trees, shrubs and grasses, meadow steppe shrubs and grasses, typical steppe shrubs and grasses, the meadow steppe and typical steppe grasses, showed higher values of physiological variables than the forest or the desert species. But for the two reproductive plant functional types (PFTs), clonal species had higher physiological variables, with averages of 22%, 15%, 23% and 14% higher than the non-clonal ones for P n, E, gs, and WUE, respectively. Such differences indicated that clonal species might have advantages over non-clonal species in utilizing environmental resources such as light, CO2, and especially water.     

Functional traits,spatial patterns and species associations: what is their combined role in the assembly of wetland plant communities?

Studies of community assembly focus on finding rules that predict which species can become member of a plant community. Within a community, species can be categorized in two ways: functional groups classify species according to their functional traits, whereas generalized guilds group species based on their (co-)occurrence, spatial distribution and abundance patterns. This study searches for community assembly rules by testing for coherence among functional groups and generalized guilds, as well as for correlations between the individual functional traits and assembly features, in two wetland plant communities in South Africa. The classifications of functional groups and generalized guilds were not consistent. However, rhizome internode length was related to fine-scale spatial pattern, suggesting that in systems dominated by clonal species (including wetlands, where recruitment sites are strongly limited) community assembly may be strongly linked to colonization ability. Functional groups do not predict guilds in wetland plant communities, precluding their use as the basis for assembly rules. However, an explicit consideration of clonal strategies and their effect on species’ spatial patterns appears to be important for understanding community assembly in systems dominated by clonal plants.     

克隆植物芦苇叶片功能性状对异质环境的响应

芦苇叶片功能性状的空间变化反映克隆植物的资源分配格局,而其与土壤环境因子的耦合关系体现了克隆植物对异质环境的生态适应策略。本研究以中国西北内陆湿地克隆植物芦苇为对象,分析了湿生生境、盐沼生境、荒漠生境条件下芦苇叶片功能性状及其对土壤环境因子的响应。结果表明: 从湿生生境到荒漠生境,芦苇叶片C、N、P含量分别下降7.2%、40.0%、64.1%,N、P利用效率增加,芦苇叶长、叶宽、叶面积、叶干重、比叶面积和叶厚度均表现出减小趋势。芦苇叶片功能性状间存在协同变化的特征,比叶面积与叶片营养元素表现出显著相关关系。土壤容重、盐分和水分分别是驱动湿生生境、盐沼生境和荒漠生境芦苇叶片功能性状变异的最重要的环境因子。     

Maximizing gain per effort by using clonal replicates in genetic tests

Summary Models for predicting cummulative genetic gain from recurrent selection applicable to predominantly outcrossing plant species are derived to include the effect of observations on clonal replicates (ramets) in addition to observations on individuals and family means. Such models are discussed with special reference to forest trees. The consequence of redistributing effort from individuals to ramets is investigated for several conditions with a fixed number of families and fixed total test size. Factors that affect the distribution of variance among sources and factors that affect individual selection intensity are the primary determinants of the optimum distribution of effort. The optimum number of ramets ranged from 1 to 6 for the conditions tested and the efficiency of redistribution (ratio of gain for the optimum distribution to the gain for the single-ramet, or non-clonal case) ranged from 1.00 to 1.20. Using clonal replicates in genetic tests usually results in increased cummulative genetic gain relative to non-clonal tests, without an increase in test effort.     

Response of plant functional traits to species origin and adaptive reproduction in weeds

Exotic species and clonal species are more competitive than co-occurring species, and several plant traits have already been identified to contribute to their superior competitiveness. In this study, 13 functional traits of 36 weed species were analyzed to determine if there was generalizable difference among these species. The results indicated that species origin significantly affected fluctuating asymmetry of width (FAW), specific leaf area (SLA), and total dry mass (TDM), with exotic species showing a higher FAW and TDM but a lower SLA. Adaptive reproduction had a significant effect so that FAW and FAA tended to be higher for non-clonal species, but RMF and R/S ratio tended to be higher for clonal species. Meanwhile, the response of these traits to species origin and reproduction was altered by their interactive effect. Detrended correspondence analysis (DCA) ordination indicated an extremely positive relationship with all leaf traits and TDM, in exotic species, but a similar relationship with SLA, TDM, and S_lw in clonal species. Clonal invasive species tended to have higher TDM, while non-clonal invasive species tended to have higher FA.     

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.     

嘉峪关草湖湿地植物功能群组成及其性状对不同生境的响应

植物功能群与生境的关系是生态学研究的重要内容,对于揭示植物的环境适应机制、群落构建机制、预测植被变化等具有重要意义。为研究局域尺度上干旱区湿地、沙漠、戈壁等生态系统功能群的环境适应机制,在嘉峪关草湖国家湿地公园内,基于植物功能性状数据,研究了荒漠区不同生境条件下的功能群组成和功能性状特性。结果表明:随着生境条件的改变,土壤含水量、含盐量、容重、pH值等土壤性质发生显著变化;植物的功能群组成和功能性状会发生适应性的变化,沼泽湿地生境内植被的功能性状以克隆、C4光合途径、根茎型等特征为主,盐沼湿地以叶片被蜡、叶茎肉质、CS策略等特征为主,沙丘生境以叶片具毛、风播、具增厚或增长的根、一年生、CR策略等特征为主,荒漠戈壁以叶片少、具增厚或增长的根、虫媒、地上芽等特征为主。局域尺度上,生境异质性尤其是土壤条件的异质性是影响植物群落组成和植物功能性状变化的重要环境因子。     

北方典型沼泽湿地高低土壤水分下植物群落特征

北方沼泽湿地在水源供给、缓解水土流失、遏制草地沙化等方面具有重要作用,明确其植物群落物种组成和多样性特征对提升其生态系统服务功能具有重要意义。目前,在北方地区开展大尺度湿地植被调查的研究仍相对较少。土壤水分是驱动植物群落发展的主导环境因素之一,为了解高低土壤水分背景下湿地植物群落特征差异及关键驱动要素,对我国7个北方典型沼泽湿地的植物群落物种组成及多样性特征进行了调查,分析了植物群落物种组成及多样性特征与环境因子的关系,以及沼泽湿地植物群落内克隆植物的分布特征。研究结果发现不同沼泽湿地的植物群落物种组成和多样性差异显著,但无明显的地带性分布规律,物种分布呈现区域性。群落物种多样性受降水、温度、土壤养分等多种环境因素的共同影响。沼泽湿地高低土壤水分背景下植物群落的物种组成和多样性差异显著,低土壤水分下植物群落物种多样性指数显著高于高土壤水分下植物群落。低土壤水分下物种多样性主要受降水和总氮影响,而高土壤水分下物种多样性主要受温度和总磷的影响。高土壤水分下克隆植物物种数和盖度在沼泽湿地植物中占有较高的比例,表明克隆植物比非克隆植物更适应高土壤水分环境。研究结果表明了7个沼泽湿地植被的区域性...     

Are clonal plants more tolerant to grazing than co-occurring non-clonal plants in inland dunes?

Clonal traits such as clonal integration and storage functions of rhizomes or stolons may provide clonal plants with additional advantages against grazing over non-clonal plants. Here, we hypothesize that clonal species have a larger capacity for compensatory growth than co-occurring non-clonal species. In inland dunes in northern China, individual plants of two rhizomatous clonal species (Bromus ircutensis and Psammochloa villosa) and two non-clonal ones (Artemisia intramongolica and Astragalus melilotoides) were subjected to 0% (control), 50% (moderate) and 90% (heavy) shoot removal. Compared with control, heavy clipping greatly increased the relative growth rate in Bromus and Psammochloa, but decreased that in Artemisia and Astragalus. Heavy clipping affected above-ground dry weight and the number of modules more negatively in Artemisia and Astragalus than in Bromus and Psammochloa. These results support the hypothesis and suggest that clonal species are more tolerant to grazing than co-occurring non-clonal species in inland dunes.     

Clonal Growth Forms in Eastern Ladakh, Western Himalayas: Classification and Habitat Preferences

Earlier observations that plant clonality, i.e., production of potentially independent offspring by vegetative growth, increase in importance in cold climates such as in arctic and alpine regions, have been recently questioned. However, lack of data obtained using a comparable methodology throughout different regions limit such comparisons. Here we present a classification of clonal growth forms for vascular plants from East Ladakh (an arid mountain range in NW Himalaya, India), and assess the relationship of these forms with multiple environmental gradients. Based on field assessment of clonality in 540 species we distinguished 20 growth forms, which were then grouped into four broader space occupancy strategies. Occurrence in communities and relationship with environmental characteristics and altitude were analyzed using multivariate methods. The most abundant growth form was represented by non-clonal perennial species with a pleiocorm having short branches, prevailing in steppes, Caragana shrubs and screes. The most abundant clonal species were those with very short epigeogenous rhizomes, such as turf graminoids prevailing in wet Kobresia grasslands. Two principal environmental gradients, together with several abiotic variables, affected space occupancy strategies: moisture and altitude. Non-spreading integrators prevailed on shaded rocky slopes, non-spreading splitters in wet grasslands and spreading splitters at the wettest sites. Spreading integrators were the least frequent strategy predominantly occurring at the most elevated sites. Because relevance of clonality decreased with altitude and different communities host different sets of clonal growth strategies, comparison with other cold climate regions should take multiple environmental gradients into account.     

Effects of the loss of clonal integration on four sedges that differ in ramet aggregation

Although clonal growth is a dominant mode of plant growth in wetlands, the importance of clonal integration, resource sharing among ramets, to individual ramet generations (mother and daughter) and entire clones of coexisting species has not been well investigated. This study evaluated the significance of clonal integration in four sedge species of varying ramet aggregations, from clump-forming species (Clumpers –Carex sterilis, Eleocharis rostellata), with tightly aggregated ramets (rhizomes<1cm), to runner species (Runners –Schoenoplectus acutus, Cladium mariscoides), with loosely aggregated ramets. We manipulated clonal integration by either severing connections between target mother and daughter ramets or leaving connections intact, and then planted them in an intact neighborhood of a fen in Michigan, USA. We measured growth parameters of original and newly produced ramets over two growing seasons and conducted a final biomass harvest, to address four hypotheses. First, we expected integrated clones to accumulate more biomass than severed clones. However, final clone-level biomass and ramet production were the same for both treatments in all species although severing initially stimulated ramet production by Schoenoplectus and produced a more compact ramet aggregation in Cladium. Second, we hypothesized that mother ramets would experience a cost of integration, through reduced ramet or biomass production, while daughters would experience a benefit, through increased resource availability from mothers. Mother ramets of Cladium suffered a cost from integration, while Schoenoplectus mothers suffered a slight cost and Carex daughters saw a slight benefit. Finally, we hypothesized that integration would be more active in runner species than in clumper species. Indeed, we documented more active integration in runners than clumpers, but none of the study species were dependent upon integration for growth or survival once daughter ramets were established with their own roots and shoots. This study demonstrates that integration between established ramets may not be the most important advantage to clonal growth in this wetland field site. The loss of integration elicited varied responses among coexisting species in their natural habitat, somewhat but not completely related to their growth form, suggesting that a combination of plant life history traits contributes to the dependence upon clonal integration among established ramets of clonal species.     

Diversity and Frequency of Clonal Traits Along Natural and Land-Use Gradients in Grasslands of the Swiss Alps

The frequency of clonal plants in different vegetation types is known to be influenced by environmental and land-use factors. However, the underlying behavior of individual clonal traits or clonal trait diversity has received little attention. Here, we assess for species- and trait-diverse grasslands of the Swiss Alps the relative importance of temperature, soil moisture, land use and species richness on the diversity and frequency of individual compared with all clonal traits. We further analyzed how cover-weighted data alters the relationships found with commonly used presence-absence data. We combined species compositional, land-use and environmental data from 236 28-m² grassland plots with clonal trait information for 527 species following the Clonal Growth Organ (CGO) classification. Test results are based on linear models, ANOVAs and ANCOVAs. The grassland sites were 84% dominated by clonal species. Drought-prone grasslands harbored the least clonal species. No increase in clonality was detected with decreasing temperature (= altitude). Mown or pastured grasslands had more clonal species than fallows. Certain sets of traits were correlated. Rhizomatous species especially reacted strongly to climatic and land-use gradients and had highest frequencies in cold, moist and disturbed sites. Clonal diversity was strongly dependent on species richness. Cover-weighted and presence-absence based estimates were largely similar. Overall, our data outlined that common clonal traits react differently to natural and land-use gradients as well as differently to the sum of clonal traits. Also, soil moisture was more decisive than temperature (= altitude) for the presence of clonal species. Lastly, the strong correlation between species-richness and clonal trait diversity needs to be accounted for when interpreting the functional role of clonal traits.     

异质性水分环境中克隆整合对活血丹生物量分配及叶片结构特征的影响

喀斯特石漠化环境有着高度的生境异质性,异质性生境中土被不连续,土壤瘠薄,岩溶漏斗上的土壤保水性差,严重制约着喀斯特植被的生长及分布。为探究克隆植物在喀斯特地区的适应策略,本研究以喀斯特黄色石灰土为基质,选用克隆植物活血丹（Glechoma longituba）,以一个节间连接的两个分株为材料,保持节间连接或切断,种植于相邻花盆中,并施以不同浇水量,以明确不同水分可用性水平下克隆整合对活血丹生物量积累、生物量分配、叶片气孔及叶片组织特征的影响。结果显示,克隆整合显著促进活血丹生物量的积累及对根、叶的生物量分配;增加了活血丹叶气孔导度,降低了气孔指数;叶海绵组织受克隆整合影响较小,但栅栏组织及栅海比（栅栏组织/海绵组织）表现为非整合分株高于整合分株。本研究表明,克隆整合可增加活血丹胁迫分株对根、叶的投资,并以更佳的叶气孔、组织适应策略提高其在喀斯特生境中的生存与适应。     

Evolutionary and organismic constraints on the relationship between spacer length and environmental conditions in clonal plants

The rate of lateral spread of clonal plants is considered a plastic trait that responds directly to environmental conditions. However, this response is likely constrained by evolutionary history and other species attributes. Here we ask how the relationships between lateral spread and the distribution of herbaceous plant species in different environments are influenced by the type of spacer (epigeogenous versus hypogeogenous rhizome), its persistence (integrator versus splitter), and the height and phylogeny of the plant. Analysis of spacer length of 367 species from temperate to arctic Europe revealed that other plant characteristics modulate the relationship between spacer length and environmental conditions. While plants with epigeogenous rhizomes, which have usually shorter spacers than hypogeogenous rhizomes, were associated with illuminated habitats, plants with hypogeogenous rhizomes were associated with warm habitats. These relationships were also shown within the specific rhizome type, as within the group of species with epigeogenous rhizomes, those having short spacers were associated with more light. The trend toward long spacers in warm environment was detected within both epigeogenous and hypogeogenous rhizome bearing species. Splitters were found to be associated with wet, nutrient rich habitats irrespective of rhizome type. When plant height was accounted for by using the ratio of spacer length to plant height (L:H) instead of absolute spacer length, no relationship with environmental variables was found until phylogenetic relationships among the species were taken into account. This implies that overall variability in L:H ratio in the dataset masked trends common for different taxonomic groups. Future comparative studies relating particular clonal growth forms or individual traits to environmental conditions should consider the constraints arising from other growth characteristics and evolutionary history of the subject species.     

Physiological integration in an introduced, invasive plant increases its spread into experimental communities and modifies their structure

What determines the invasiveness of introduced plants is still poorly known. Many of the most invasive plant species are clonal, and physiological integration between connected individuals (ramets) of clonal plants may contribute to their ability to spread into communities and reduce performance of existing species. This contribution of integration to the invasiveness of clonal plants may be greater in denser communities. A greenhouse study was conducted to test these two hypotheses. High- and low-density communities were created by sowing seeds of eight grassland species. Each community was planted with three ramets of the stoloniferous, introduced plant Alternanthera philoxeroides that were disconnected from or left connected to ramets growing on bare soil. Connection increased the spread of Alternanthera within a community, but did not reduce community biomass. Alternanthera grew less in high-density communities, but connection did not improve its growth more than in low-density communities. Low-density communities had higher evenness when Alternanthera was connected than when it was disconnected because shoot mass was lower in the more abundant species in the community and higher in the less abundant ones. These results partly supported the first hypothesis, but not the second. The effect of integration on community structure could be due to higher resource import by the ramets of Alternanthera closer to the dominant species. Integration therefore can increase the initial spread of new clonal plant species into communities and modify the effects of this spread on community structure.     

Differential influence of clonal integration on morphological and growth responses to light in two invasive herbs

Background and aims

In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions.

Methods

In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85% shade and their morphological and growth responses were assessed.

Key results

The influence of clonal integration on the light reaction norm (connection×light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection×light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mother plants invest in daughter ramets prior to their own branches.

Conclusions

Clonal integration modifies light reaction norms of morphological and growth traits in a species-specific manner for A. philoxeroides and P. canescens, but it improves the establishment of juvenile ramets of both species in light-limiting environments by promoting their growth during early development. This factor may be partially responsible for their ability to successfully colonize native plant communities.