Fitness and evolution in clonal plants: the impact of clonal growth

Seeds have often been emphasized in estimates of plant fitness because they are the units that carry genes to the next generation, disperse, and found new populations. We contend that clonal growth also needs to be considered when estimating fitness in clonal plants, regardless of whether fitness is measured from a genet or ramet perspective. Clonal growth affects genet fitness through both genet persistence and seed production. It affects ramet fitness through new ramet production, because both seeds and clonal propagants are considered offspring. The differential production of clonal propagants will contribute to fitness differences among individuals which may result in population-level changes in allele frequencies (i.e. microevolution). We describe a form of selection unique to clonal organisms, genotypic selection, that can result in evolution. Genotypic selection occurs when genotypically based traits are associated with differences in the rate of ramet production. It can lead to evolutionary change in quantitative trait means both directly and indirectly. It leads directly to change in the ramet population by increasing the proportion of ramets with more advantageous trait values. From the genet perspective, it leads indirectly to evolution within and among populations whenever significant portions of the genetic effect on a trait are inherited through seed. We argue that under most conditions, clonal growth will play a major role in the microevolution of clonal plants.     

A simple model for the influence of habitat resource availability on lateral clonal spread

Plant clonal spread is ubiquitous and of great interest, owing both to its key role in plant community assembly and its suitability for plant behaviour research. However, mechanisms that govern spreading distance are not well known. Here we link spacer costs and below-ground competition in a simple model of growth in a homogeneous below-ground environment, in which optimal distance between ramets is based on minimizing the sum of these costs. Using this model, we predict a high prevalence of clonal growth that does not employ spacers in resource-poor environments and a nonlinear increase in spreading distance in response to increasing below-ground resource availability. Analysis of database data on clonal growth in relationship to below-ground resource availability revealed that patterns of the spread based on stolons is compatible with the model''s predictions. As expected, model prediction failed for rhizomatous species, where spacer sizes are likely to be selected mainly to play roles other than spread. The model''s simplicity makes it useful as a null model in testing hypotheses about the effects of environmental heterogeneity on clonal spread.     

Restoration of three forest herbs in the Liliaceae family by manipulating deer herbivory and overstorey and understorey vegetation

Background: Research on herbaceous vegetation restoration in forests characterised by overstorey tree harvests, excessive deer herbivory, and a dominant fern understorey is lacking. Most of the plant diversity found in Eastern hardwood forests in the United States is found in the herbaceous understorey layer. Loss of forest herbaceous species is an indicator of declining forest conditions.

Aims: The combined effects of deer herbivory, competitive understorey vegetation removal, and overstorey tree removal on the abundance and reproductive capacity of three understorey herbs in the Liliaceae family were evaluated.

Methods: A split-plot randomised block design was used with three replicates. Treatments included three harvest intensities, fenced/unfenced, herbicide/no herbicide-treated, prescribed burn/no prescribed burn, and all combinations. A generalised linear model was used to compare treatment effects over 8 years.

Results: Both fruit production and cover increased significantly in fenced areas for all three species. There was a significant 6-year recovery period for cover of the three species in response to herbicide. There was a significant 4-year recovery period of fire-treated plots for fruit production of the three species. The most intensively cut, fenced, and herbicide-treated plots had the greatest increases in sapling and Rubus spp. cover. Cover and fruit production of the three herbs were significantly greatest in the moderate-cut treatment.

Conclusions: Restoration of these three liliaceous species is most likely to occur in Eastern deciduous forests and similar forests using a combined fenced and moderate-cut treatment.     

Using simulated herbivory to predict the efficacy of a biocontrol agent: the effect of manual defoliation and Macaria pallidata Warren (Lepidoptera: Geometridae) herbivory on Mimosa pigra seedlings

Abstract  The effect of manual defoliation, and real and simulated herbivory by the leaf-feeding moth Macaria pallidata on the growth and biomass of Mimosa pigra seedlings was assessed to determine the likely success of defoliating herbivores as biocontrol agents against the weed. Manual removal of at least half of all leaf material, maintained over a 6-week period, significantly suppressed plant growth, as did a single week of herbivory by M. pallidata larvae. This suggests that M. pallidata and other defoliators have the potential to be highly successful agents in the biological control program for M. pigra.     

Herbivory and ramet performance in the clonal herb Trientalis europaea L.

1 The intensity of leaf damage caused by invertebrate herbivores and grazing by vertebrates and their effects on clonal growth, survival and reproduction were examined in a population of the forest herb Trientalis europaea during 4 years.
2 Levels of herbivory were low and varied between years in the studied population. Larger ramets were exposed to a greater risk of both grazing and leaf defoliation. Ramets that suffered leaf damage in 1 year experienced an increased probability of leaf damage in succeeding years. The probability of suffering herbivore attack was independent among the ramets belonging to the same clonal fragment, suggesting that clonal propagation might operate as a risk-spreading strategy in this species.
3 Leaf damage did not affect any measure of plant performance, probably due to the low amounts of leaf area removed by invertebrate herbivores. In contrast, vertebrate grazing affected all phases of the pseudo-annual life cycle of T. europaea . Grazing prevented flowering and fruiting, increased ramet mortality during summer and decreased tuber production. Furthermore, grazed ramets produced shorter stolons and smaller tubers, which in turn had a lower winter survival and produced smaller ramets in the following growing season. The large impact of grazing was due to the consumption of the whole of the single shoot of ramets of T. europaea . Although regrowth was possible, secondary shoots were significantly smaller and assimilation was delayed.
4 Tubers originating from grazed ramets were placed shallower than tubers from ungrazed ramets. This could be due to an alteration in the growth pattern of stolons of the grazed ramets.     

The relative contributions of sexual reproduction and clonal propagation in Opuntia rastrera from two habitats in the Chihuahuan Desert

1 The clonal cactus Opuntia rastrera shows predominantly sexual reproduction in grasslands (GH) and clonal propagation in nopaleras (NH). We assessed the effects of light, herbivory, water availability and the habitat an offspring came from on the survival and growth of sexual or clonal offspring (i.e. seedlings and cladodes), through 3- and 4-year common garden and short-term greenhouse experiments.
2 Shading by nurse plants increased seedling survival in the field by an order of magnitude, and a small additional advantage due to predator protection by grasses was observed. Strong herbivory transforms a facultative nurse–protégé relationship for seedlings into an obligatory one.
3 In the greenhouse seedlings grew better under shade, but in the field the production of the first cladode was delayed in seedlings in the more shaded GH. Competition for soil resources may be more intense under a dense grass tussock than under a open shrub, thus affecting the nurse–protégé relationship. Seedling survival under nurse plants was similar in GH and NH, but higher plant cover suggests that a larger number of seedlings will establish in GH in the long term.
4 Cladode survival was higher in NH. Cladodes were more successful than seedlings at establishing in intercanopy areas, possibly due to physiological differences as well as their ability to survive partial predation. Cladode survival in intercanopy areas may explain the enhanced clonal propagation in the more open NH scrubland, together with their susceptibility to the flooding which affects GH.
5 The high seedling and cladode survival in the greenhouse experiments contrasted with that observed in the field, indicating that survival is determined by the interaction between herbivores, plants and abiotic conditions rather than the physiological aptitude of the plants.     

Effect of the specialist herbivore Luehdorfia puziloi on the performance of a woodland herbaceous plant, Asarum heterotropoides

Plants show many responses to herbivore damage caused by insects. We investigated the effect of the specialist leaf herbivore Luehdorfia puziloi on the performance of a deciduous forest understory perennial herb Asarum heterotropoides . We conducted artificial defoliation experiments with different levels of damage (0, 50 or 100% clipping) in the early growing period (flowering time: early May) and in the late growing period (when natural herbivory by L. puziloi larvae occurs: early June) in a natural population. Effects of the defoliation treatments on reproduction (seed-set ratio) of A. heterotropoides in the treated year and on survival and development of the plant in the subsequent year were investigated. Severe defoliation conducted in the early period resulted in less current-year seed production and increased regression to a dormant or non-reproductive stage in the following year, which would reduce the probability of future flowering. In contrast, defoliation conducted in the late period had no effect on the reproduction, survival and development of A. heterotropoides . Similarly, we found no obvious correlation between the damaged leaf area during the later period and seed set. Our results show that the specialist herbivore L. puziloi did not influence the performance of A. heterotropoides by damaging the plant later in the season when it was tolerant of damage.     

Timing of disturbance changes the balance between growth and survival of parent and offspring ramets in the clonal forest understory herb Uvularia perfoliata

Preformation of organs involves the initiation of vegetative and generative tissues at least one season before they are actually produced. It is a strategy to deal with environments characterized by predictable seasonality as it enables fast growth of plants at the onset of favorable conditions. However, early preformation also strongly restricts plants in their response to unpredictable environmental changes and disturbance. In this study we investigated the response of the clonal forest understory herb Uvularia perfoliata to disturbance and resource limitation. In U. perfoliata shoot characteristics, as well as vegetative and sexual reproduction are determined at the end of the previous growing season. Plants were grown under two light levels and the rhizome connection between parent and offspring ramets were severed at various times during the growing period. Disturbance did not affect total biomass accumulation but it did affect the relative allocation and survival probability of parents and offspring ramets. Early severing resulted in increased survival chance and future fitness of the parent ramet, while late severing resulted in a higher survival chance and increased fitness of offspring ramets. The response was mediated by plant size and resource availability. These results show that the life history of U. perfoliata includes the possibility to alleviate the effects of disturbance even though the species is characterized by strong developmental canalization through organ preformation.Co-ordinating editor: J. Tuomi     

Florivory: the intersection of pollination and herbivory

Plants interact with many visitors who consume a variety of plant tissues. While the consequences of herbivory to leaves and shoots are well known, the implications of florivory, the consumption of flowers prior to seed coat formation, have received less attention. Herbivory and florivory can yield different plant, population and community outcomes; thus, it is critical to distinguish between these two types of consumption. Here, we consider the ecological and evolutionary consequences of florivory. A growing number of studies recognize that florivory is common in natural systems and in some cases surpasses leaf herbivory in magnitude and impact. Florivores can affect male and female plant fitness via direct trophic effects and through altered pathways of species interactions. In particular, florivory can affect pollination and have consequences for plant mating and floral sexual system evolution. Plants are not defenceless against florivore damage. Concepts of resistance and tolerance can be applied to plant–florivore interactions. Moreover, extant theories of plant chemical defence, including optimal defence theory, growth rate hypothesis and growth differentiation–balance hypothesis, can be used to make testable predictions about when and how plants should defend flowers against florivores. The majority of the predictions remain untested, but they provide a theoretical foundation on which to base future experiments. The approaches to studying florivory that we outline may yield novel insights into floral and defence traits not illuminated by studies of pollination or herbivory alone.     

高寒草甸退化对短穗兔耳草克隆生长特征的影响

以调查统计和比较样地法研究了江河源区高寒草甸退化对典型匍匐茎植物短穗兔耳草克隆生长特征的影响。结果表明,退化草甸的植物群落结构、功能以及土壤特征发生了明显地变化,继而对短穗兔耳草无性系的克隆生长行为和形态特性产生了影响。高寒草甸退化后短穗兔耳草的匍匐茎有所增多,分支强度加大。退化草甸内短穗兔耳草的基株高度小于未退化草甸,根长大于未退化草甸,基株的叶片数目间没有明显差别。退化草甸内短穗兔耳草的分株高度显著小于未退化草甸,分株叶数明显多于未退化草甸,而根长尽管大于未退化草甸,但差异不显著。短穗兔耳草匍匐茎长度在未退化草甸内明显大于退化草甸,匍匐茎茎生叶数和匍匐茎粗度也在未退化草甸大于退化草甸。短穗兔耳草在未退化草甸用于克隆繁殖的能量投资比例高于退化草甸,其中未退化草甸内短穗兔耳草基株的干重比例略低于退化草甸,分株和匍匐茎的干重比例高于退化草甸。高寒草甸退化对短穗兔耳草克隆生长特征的这些影响,是其对高寒草甸退化导致的资源和生境差异的反应,也是对资源利用达到的最合理状态,是一种选择适应的结果。     

Importance of clonal plants and plant species diversity in the Northeast China Transect

In plant communities, the internal (genet-level) control mechanisms on a spatio-temporal scale of clonal plants impose strong constraints on spatial pattern as well as on competitive relations and, thus, species coexistence. Therefore, the presence of clonal species within a plant community affects spatio-temporal dynamics and plant species diversity. We examined the distribution of plants with different clonal growth forms in the Northeast China Transect (NECT) and correlated plant species diversity with the importance of clonal plants, and the importance of phalanx and guerilla clonal plants. Phalanx clonal plants were more abundant in western communities where the altitude was higher and both the soil nitrogen contents and precipitation were relatively low. Whereas guerilla clonal plants were more abundant in the middle of the NECT where the precipitation, mean annual temperature and photosynthetically active radiation were relatively high. In the relatively productive temperate typical steppe, plant species diversity was negatively correlated with the importance of phalanx clonal plants and positively correlated with the importance of guerilla clonal plants. In relatively unproductive temperate desert steppe, plant species diversity was positively correlated with the importance of both phalanx and guerilla clonal plants.     

Impact of the flower smut Ustilago cynodontis (Ustilaginaceae) on the performance of the clonal grass Cynodon dactylon (Gramineae)

Effects of the flower-infecting smut fungus Ustilago cynodontis on the growth, survival, and competitiveness of the clonal rhizomatous perennial grass Cynodon dactylon were explored through a series of glasshouse experiments. Ustilago cynodontis sterilized C. dactylon by replacing floral structures with a teliospore-producing fungal stroma. Infection caused a reduction in overall dry matter production and the growth rate of stolons, changed the allocation of resources between roots and shoots, and affected the survival of C. dactylon plants growing in pots under crowded conditions in a naturally lit glasshouse. In mixtures of healthy and infected plants, the growth of infected ones was disproportionally affected. Infection had no effect on the germination or emergence of seedlings. Transmission of the pathogen along stolons was incomplete, especially as the length of the stolon increased. The nutrient status of the soil had no significant effect on stolon growth. Results are discussed in terms of the differential effect of systemic floral smut diseases on tiller-forming and clonal rhizomatous perennial grasses.     

川西北高寒沙区切断根茎对赖草和沙生苔草克隆生长的影响

为了研究高原亚寒带沙化生境中切断根茎对克隆植物基株扩展能力和分株定居能力的影响,在川西北若尔盖高原沙化区内,对根茎禾草赖草和沙生苔草进行了以切断根茎为处理的野外实验。结果表明,赖草和沙生苔草基株的幼小部分(观测单元)地上枝总长度增量、主根茎长度增量和根茎总长度增量显著减少,而对根茎数增量、主根茎节增量和根茎节总数增量影响不显著;赖草观测单元地上枝数增量显著减少,而对沙生苔草地上枝数增量无影响;赖草地上枝与根茎的相关性质发生逆转。这表明．在高原亚寒带半湿润沙化生境中．克隆整合效应显著促进基株幼小部分地上枝和根茎的伸长生长,但对新生根茎的产生和根茎节分化没有影响;切断根茎处理导致赖草、沙生苔草生殖生长与营养生长间竞争加剧,同时使赖草地上部分与地下部分间竞争加剧;观测单元在缺少与基株(或上级株系)的克隆整合作用时,赖草受到的影响大于沙生苔草。     

The influence of below ground herbivory and plant competition on growth and biomass allocation of purple loosestrife

Experiments investigating plant-herbivore interactions have primarily focused on above-ground herbivory, with occasional studies evaluating the effect of below-ground herbivores on plant performance. This study investigated the growth of the wetland perennial Lythrum salicaria (purple loosestrife) under three levels of root herbivory by the weevil Hylobiustransversovittatus and three levels of plant competition by the grass Phleumpratense in a common garden. Plant growth, flowering phenology, and biomass allocation patterns of purple loosestrife were recorded for two growing seasons. During the first year, root herbivory reduced plant height; plant competition delayed flowering; and the interaction of root herbivory and plant competition resulted in reductions in plant height, shoot weight and total dry biomass. Plant competition or larval feeding did not affect the biomass allocation pattern in the first year. These results indicate the importance of interactions of plant competition and herbivory in reducing plant performance – at least during the establishment period of purple loosestrife. In the second growing season, root herbivory reduced plant height, biomass of all plant parts, delayed and shortened the flowering period, and changed the biomass allocation patterns. Plant competition delayed flowering and reduced the dry weight of fine roots. The interaction of root herbivory and plant competition delayed flowering. Root herbivory was more important than plant competition in reducing the performance of established purple loosestrife plants. This was due, in part, to intense intraspecific competition among the grass individuals effectively preventing shoot elongation of P. pratense and resulting in a carpet like growth. Received: 3 April 1997 / Accepted: 27 July 1997     

Circumnutation behavior of an exotic honeysuckle vine and its native congener: influence on clonal mobility

Virtually all plant parts rotate slightly about a central axis, a movement called circumnutation, but vines show exaggerated circumnutation. This study contrasts circumnutation in two congeneric twining vines, specifically focusing on differences in erect and prostrate shoots, and examines the impact of circumnutation on exploitation of available climbing supports and exploration for more distant ones. Elongating shoots of Lonicera japonica and L. sempervirens growing in a common garden were classed as (1) erect but not climbing, (2) climbing on a trellis, or (3) prostrate, and their circumnutation quantified by tracking the compass direction of shoot tips. To quantify the impact of different circumnutation behaviors, the climbing success of erect shoots and the maximum dispersion and rooting success of prostrate shoots were measured. Erect shoots of both species circumnutated at similar rates (averaging 31°/h), and did not differ in their success rate of exploiting trellises (averaging 76.8%). Prostrate shoots differed, with those produced by L. japonica having reduced circumnutation. In contrast, prostrate shoots of L. sempervirens continued to circumnutate as much as erect shoots. The specialized circumnutation behavior of the prostrate shoots of L. japonica results in increased rooting success and maximum dispersion compared to the unspecialized shoots of L. sempervirens.     

Genetic variation in clonal traits of Trifolium repens and species interactions

Plants exhibit a great variety of types of clonal growth. Moderate variation in clonal traits often exists even within species. The consequences of these variations for species interaction are of great interests to ecologists. In this paper, I address the small-leaved (phalanx) to large-leaved (guerrilla) variation in white clover ( Trifolium repens ), and discuss its consequences for species and genotype coexistence. I also address the clonal and sexual resource allocation variants within the large-leaved type. Small-leaved and large-leaved genotypes differ in various aspects of clonal growth. The large-leaved genotype displays greater phenotypic plasticity but is less physiologically integrated than the small-leaved genotype. We examined the consequences in a grazed sward, where white clover and zoysia grass coexist. In this sward, white clover is patchily distributed. We first tested the hypothesis that the large-leaved genotype is more advantageous in growth than the small-leaved genotype. Results from both common garden and competition experiments supported the hypothesis. Second, we tested the hypothesis that within large-leaved plants, the clonal subtype (which invests more resources to stolons but less in flower heads than the sexual one) is more advantageous than the sexual one because it is more competitive. This hypothesis was rejected. Both subtypes coexisted in the sward. This is probably because the sexual subtype is superior for interpatch migration than the clonal one. Both subtypes differ in advantages they offer for between-patch and within-patch processes, which promotes their coexistence. Finally, field monitoring of the behavior of a large-leaved clone is described. This monitoring was conducted in a moderately grazed sward, where microenvironmental heterogeneity is extremely high in time and space.     

Tolerance of Gentianella campestris in relation to damage intensity: an interplay between apical dominance and herbivory

Meristem allocation models suggest that the patterns of compensatory regrowth responses following grazing vary, depending on (i) the number of latent meristems that escape from being damaged, and (ii) the activation sensitivity of the meristems in relation to the degree of damage. We examined the shape of compensatory responses in two late-flowering populations (59°20′N and 65°45′N) of the field gentian. Plants of equal initial sizes were randomly assigned to four treatment groups with 0, 10, 50 and 75% removal of the main stalk. The plants were clipped before flowering, and their performance was studied at the end of the growing season. The northern population showed a linear decrease in shoot biomass and fecundity with increasing biomass removal, while the response in the southern population was quadratic with maximum performance at the damage level of 50% clipping. This nonlinear shape depended upon the activation sensitivity of dormant meristems in relation to their position along the main stem. The highest plant performance was achieved by inflicting intermediate damage which induced regrowth from basally located meristems. In contrast, the topmost branches took over the dominance role of the main stem after minor apical damage (10% clipping). Consequently, the breakage of apical dominance is a necessary precondition of vigorous regrowth in this species. However, compensation in the field gentian is unlikely to be a mere incidental by-product of apical dominance. The ability to regrow from basally located meristems that escape from being damaged by grazing may well be a sign of adaptation to moderate levels of shoot damage. This revised version was published online in July 2006 with corrections to the Cover Date.     

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.Co-ordinating editor: J. Tuomi