Patterns of Differentiation in Wiregrass (Aristida beyrichiana): Implications for Restoration Efforts

Aristida beyrichiana (wiregrass) is increasingly being planted in restoration projects across the southeastern coastal plain, with little focus on genetic differences among populations across the region. Local and regional population differentiation for establishment and growth traits were examined in common garden and reciprocal transplant experiments. Seeds from up to 20 plants from each of seven populations were collected in northern and central Florida sites that encompassed gradients of soils, hydrology, and temperature. Reciprocal seed transplants using three of the common garden populations were conducted in two consecutive years. In the common garden, significant population differences were seen in seed weight, seedling emergence and survival, tiller height, number of tillers, the relationship between tiller number and tiller height, and flowering. Variation among maternal families was seen in tiller number and in the relationship between tiller number and tiller height. The reciprocal transplant study did not detect either local adaptation to sites of origin or consistent superiority of one source population or planting site in seedling establishment. These results suggest that the probability of seedling establishment is primarily dependent on environmental conditions rather than genetic differences. Genetic variation for traits related to fitness (e.g., tiller number) may be retained within populations because phenotypically plastic growth responses of seedlings to environmental variation buffer genetic variation against the action of selection. But despite the lack of evidence for genetic influences on initial establishment in wiregrass, our common garden study suggests genetic differences among populations. This result, when combined with previous results indicating local adaptation in later life stages of wiregrass, suggests that restoration efforts involving this species should use local seed sources from sites with similar soil and hydrological conditions.     

Glacial History Affected Phenotypic Differentiation in the Alpine Plant,Campanula thyrsoides

Numerous widespread Alpine plant species show molecular differentiation among populations from distinct regions. This has been explained as the result of genetic drift during glacial survival in isolated refugia along the border of the European Alps. Since genetic drift may affect molecular markers and phenotypic traits alike, we asked whether phenotypic differentiation mirrors molecular patterns among Alpine plant populations from different regions. Phenotypic traits can be under selection, so we additionally investigated whether part of the phenotypic differentiation can be explained by past selection and/or current adaptation. Using the monocarpic Campanula thyrsoides as our study species, a common garden experiment with plants from 21 populations from four phylogeographic groups located in regions across the Alps and the Jura Mountains was performed to test for differentiation in morphological and phenological traits. Past selection was investigated by comparing phenotypic differentiation among and within regions with molecular differentiation among and within regions. The common garden results indicated regional differentiation among populations for all investigated phenotypic traits, particularly in phenology. Delayed flowering in plants from the South-eastern Alps suggested adaptation to long sub-mediterranean summers and contrasted with earlier flowering of plants experiencing shorter growing seasons in regions with higher elevation to the West. Comparisons between molecular and phenotypic differentiation revealed diversifying selection among regions in height and biomass, which is consistent with adaptation to environmental conditions in glacial refugia. Within regions, past selection acted against strong diversification for most phenotypic traits, causing restricted postglacial adaptation. Evidence consistent with post-glacial adaptation was also given by negative correlation coefficients between several phenotypic traits and elevation of the population''s origin. In conclusion, our study suggests that, irrespective of adaptation of plants to their current environment, glacial history can have a strong and long-lasting influence on the phenotypic evolution of Alpine plants.     

Clinal differentiation during invasion: <Emphasis Type="Italic">Senecio inaequidens</Emphasis> (Asteraceae) along altitudinal gradients in Europe

The dynamics of plant population differentiation may be integral in predicting aspects of introduced species invasion. In the present study, we tested the hypothesis that European populations of Senecio inaequidens (Asteraceae), an invasive species with South African origins, differentiated during migration from two independent introduction sites into divergent altitudinal and climatic zones. We carried out 2 years of common garden experiments with eight populations sampled from Belgian and ten populations from French altitudinal transects. The Belgian transect followed a temperature and precipitation gradient. A temperature and summer drought gradient characterized the French transect. We evaluated differentiation and clinal variation in plants germinated from field-collected seed using the following traits: days to germination, days to flowering, height at maturity, final plant height and aboveground biomass. Results showed that S. inaequidens populations differentiated in growth traits during invasion. During the 1st year of sampling, the results indicated clinal variation for growth traits along both the Belgium and French altitudinal transects. Data from the 2nd year of study demonstrated that with increasing altitude, a reduction in three growth traits, including plant height at maturity, final plant height and aboveground biomass, was detected along the French transect, but no longer along the Belgian one. Phenological traits did not exhibit a clear clinal variation along altitudinal transects. The possible evolutionary causes for the observed differentiation are discussed. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differentiation and adaptation in <Emphasis Type="Italic">Brassica nigra</Emphasis> populations: interactions with related herbivores

Local adaptation and population differentiation of plants are well documented, but studies on interactions with natural enemies are rare. In particular, evidence for plant adaptation to the local biotic environment, such as herbivores remains poor. We used the black mustard Brassica nigra, an annual species of river valley and coastal habitats to (1) analyse population differentiation in plant traits and herbivory in a common garden experiment, (2) examine home versus away differences in a reciprocal transplant experiment and (3) test whether plants are adapted to local herbivores or vice versa under standard greenhouse conditions. In the common garden experiment, we found significant differentiation in plant traits, leaf damage and herbivore number among seven populations of B. nigra from France and Germany (distance 15–1,000 km). Differences were particularly strong among coastal and river valley populations and did not necessarily increase with geographical distance. A herbivore removal treatment did not change population differentiation when compared with the control allowing natural colonisation. The reciprocal transplant experiment at a scale of 15–30 km did not reveal local plant adaptation, whilst one dominant herbivore species (Meligethes aeneus) occurred in significantly higher numbers on local plants. A greenhouse experiment combining three aphid (Brevicoryne brassicae) and plant populations of the same provenance indicated herbivore adaptation to their local plants rather than plant adaptation, but overall contrasts between local and non-local combinations were not significant. The results suggest that herbivores may counteract local plant adaptation to other environmental factors. Our study has important implications for plant translocations in ecological restoration projects.     

Flowering requirements of Polymnia canadensis (Asteraceae) and their influence on its life history variation

The purpose of this study is to determine the flowering requirements of Polymnia canadensis and how they correspond to the occurrence of winter annuals, biennials, and short-lived monocarpic perennials in this species. Polymnia canadensis has a vernalization requirement for flowering, and even very small plants (i.e., those with one pair of leaves) can be vernalized. Vernalized plants can flower under both long- and short days. However, to flower plants must attain a minimum postvernalization size. Plants of this primarily monocarpic species that do not die after they flower once require another period of vernalization to flower a second time (i.e., to be dicarpic). Vernalized plants exposed to high temperatures can be devernalized; these must be re-vernalized in order to flower.     

Ecological life history of Polymnia canadensis,a monocarpic species of the North American Temperate Deciduous Forest: Demography

A demographic investigation was conducted to assess variation in life history of Polymnia canadensis (Asteraceae), a geographically-widespread, herbaceous species of deciduous forests in eastern North America. During 1985-1994, 23,063 seedlings of P. canadensis were monitored at five central Kentucky study sites. Numbers at the end were: biennials, 554; triennials, 142; winter annuals, 16; monocarpic perennials, 2; tricarpic perennials (three years), 3; and dicarpic perennials (some skipped years) that matured in the first year of life, 23; in the second year, 60; and in the third year, 9. Weekly cohorts of P. canadensis generally exhibited Deevey Type III survivorship with highest seedling mortality in summer associated with low soil moisture. Wide spatial variation in life history was displayed by the fact that fall germination cohorts at dry sites generally had greater germination and survivorship than at mesic sites during seedling establishment, while the reverse was true for spring cohorts. This led to more reproductive individuals in fall germination cohorts than spring cohorts at dry sites and generally the opposite case at mesic sites. Forest shade in mesic sites caused slower growth and a greater frequency of longer-lived reproductive individuals than in open, dry sites, but it also resulted in higher survivorship than at dry sites during moderate drought. Annual population growth rate averaged across four years was not significantly different between a dry site and mesic site, indicating that despite forest shade, P. canadensis persisted in the mesic site as well as it did in the dry site. Population structure varied among years, seasons, and study sites; at two study plots, a 2-year flowering cycle of mass seeding and senescence persisted for 4 years. At a smaller scale, there was little difference in survivorship between study plots within sites or between quadrats within study plots, while in a few instances there were large differences in the number of reproductive individuals. Weekly cohorts that germinated early within seasonal cohorts had greater number of reproductive individuals than later weekly cohorts, but not consistently greater survivorship. Several droughts induced temporal variation that was as important as spatial variation. During these droughts, the population size of all cohorts that germinated prior to the droughts declined to zero at all sites, and biennials were the longest lived type of reproductive individuals. During drought, Deevey Type I survivorship was prevalent, and lack of seed rain led to dependence on persistent seed banks for recruitment in some seasonal cohorts.     

Non‐native conditions favor non‐native populations of invasive plant: demographic consequences of seed size variation?

Trait differences between native and non‐native populations may explain the greater abundance and impact of some organisms in their non‐native ranges than in their native ranges. Here, we conducted reciprocal common gardens in southwestern Turkey (home) and central Argentina (away) to explore the hypothesis that the greater success of the invasive ruderal Centaurea solstitialis in Argentina than Turkey is partially explained by differences between home and away populations. Unusual among common gardens, our experimental design included seed additions to explicitly evaluate population level responses, as well as disturbance and no‐disturbance treatments. We documented seed mass in native and non‐native populations, and during the experiment, we periodically measured density, plant size, and herbivory. After six months, we determined the establishment of plants for populations from both origins in both home and away common gardens. Seed mass was two times larger for Argentinean than Turkish populations. Density, plant size and final establishment were also greater for plants from Argentinean than from Turkish populations, but only in the common garden in Argentina. In Turkey, no differences between population origins were detected for these variables. Herbivory was similar for populations from both origins in both common gardens. As expected, disturbance generally increased plant performance in both regions. Our results suggest that increased seed size in non‐native populations may have demographic consequences under non‐native conditions that can contribute to the invasive success of C. solstitialis. This is the first reciprocal common garden that supports the idea that seed size variation contributes to demographic differences for an invasive species between native and non‐native distributions, but our findings further suggest that seed size effects on demography depend on the ecological context in which population processes occur.     

Different gardens, different results: native and introduced populations exhibit contrasting phenotypes across common gardens

Invasive plants may respond through adaptive evolution and/or phenotypic plasticity to new environmental conditions where they are introduced. Although many studies have focused on evolution of invaders particularly in the context of testing the evolution of increased competitive ability (EICA) hypothesis, few consistent patterns have emerged. Many tests of the EICA hypothesis have been performed in only one environment; such assessments may be misleading if plants that perform one way at a particular site respond differently across sites. Single common garden tests ignore the potential for important contributions of both genetic and environmental factors to affect plant phenotype. Using a widespread invader in North America, Cynoglossum officinale, we established reciprocal common gardens in the native range (Europe) and introduced range (North America) to assess genetically based differences in size, fecundity, flowering phenology and threshold flowering size between native and introduced genotypes as well as the magnitude of plasticity in these traits. In addition, we grew plants at three nutrient levels in a pot experiment in one garden to test for plasticity across a different set of conditions. We did not find significant genetically based differences between native and introduced populations in the traits we measured; in our experiments, introduced populations of C. officinale were larger and more fecund, but only in common garden experiments in the native range. We found substantial population-level plasticity for size, fecundity and date of first flowering, with plants performing better in a garden in Germany than in Montana. Differentiation of native populations in the magnitude of plasticity was much stronger than that of introduced populations, suggesting an important role for founder effects. We did not detect evidence of an evolutionary change in threshold flowering size. Our study demonstrates that detecting genetically based differences in traits may require measuring plant responses to more than one environment.     

Reintroduction of Castilleja levisecta: Effects of Ecological Similarity,Source Population Genetics,and Habitat Quality

A suite of ecological and genetic factors are likely to contribute to reintroduction performance. Potential factors include the ecological similarity between seed source and introduction site, population size and genetic diversity of seed sources, and the habitat quality of the introduction site. We conducted common garden experiments with golden paintbrush (Castilleja levisecta), an endangered species from the Pacific Northwest, U.S.A., in order to test hypotheses about reintroduction performance and to provide management recommendations. Ten common gardens, each composed of C. levisecta individuals grown from seed from six of the remaining populations, were planted into field conditions and monitored during two growing seasons. Plant community characteristics were important predictors of observed variation in C. levisecta performance. Exotic species‐cover at common garden sites was associated with a reduction in performance of first‐year C. levisecta transplants, while survival to the second growing season increased with increasing similarity in plant functional groups between source and common garden sites. Although measures of genetic diversity, population size, and geographic distance are often used to make conservation decisions during species recovery, here they were poor predictors of C. levisecta performance and establishment. We recommend choosing material for reintroduction from ecologically similar habitats, rather than those most proximate geographically, and selecting recovery sites with low exotic species abundance.     

Experiments on growth interactions between two invasive macrophyte species

Abstract. The success of invasive species has been attributed to the ability to displace other species by direct competition. We studied growth and possible competition between the two macrophyte species Elodea nuttallii and E. canadensis, because the former has been observed to replace the latter in the field. Additional experiments were conducted in aquaria with mixed plantings of Elodea species. Species growth was measured and competitive abilities of each species determined by applying the reciprocal yield model to mean plant weight and length. In monocultures the growth rates of the two species were similar, while in mixtures the growth rate of E. canadensis was significantly lower than that of E. nuttallii. E. canadensis was more sensitive to intraspecific than to interspecific neighbours, whereas E. nuttallii was indifferent to the presence of neighbours. Differential growth characteristics of Elodea species can explain the displacement of E. canadensis by E. nuttallii under eutrophic field conditions.     

An explicit test for the contribution of environmental maternal effects to rapid clinal differentiation in an invasive plant

Population differentiation of alien invasive plants within their non‐native range has received increasingly more attention. Common gardens are typically used to assess the levels of genotypic differentiation among populations. However, in such experiments, environmental maternal effects can influence phenotypic variation among individuals if seed sources are collected from field populations under variable environmental regimes. In the present study, we investigated the causes of an altitudinal cline in an invasive plant. Seeds were collected from Senecio inaequidens (Asteraceae) populations along an altitudinal gradient in southern France. In addition, seeds from the same populations were generated by intra‐population crossings in a climatic chamber. The two seed lots were grown in a common garden in Central Belgium to identify any evidence of environmentally induced maternal effects and/or an altitudinal cline in a suite of life‐history traits. Results failed to detect any environmental maternal effects. However, an altitudinal cline in plant height and above‐ground biomass was found to be independent of the maternal environment.     

Trait differentiation and adaptation of plants along elevation gradients

Studies of genetic adaptation in plant populations along elevation gradients in mountains have a long history, but there has until now been neither a synthesis of how frequently plant populations exhibit adaptation to elevation nor an evaluation of how consistent underlying trait differences across species are. We reviewed studies of adaptation along elevation gradients (i) from a meta‐analysis of phenotypic differentiation of three traits (height, biomass and phenology) from plants growing in 70 common garden experiments; (ii) by testing elevation adaptation using three fitness proxies (survival, reproductive output and biomass) from 14 reciprocal transplant experiments; (iii) by qualitatively assessing information at the molecular level, from 10 genomewide surveys and candidate gene approaches. We found that plants originating from high elevations were generally shorter and produced less biomass, but phenology did not vary consistently. We found significant evidence for elevation adaptation in terms of survival and biomass, but not for reproductive output. Variation in phenotypic and fitness responses to elevation across species was not related to life history traits or to environmental conditions. Molecular studies, which have focussed mainly on loci related to plant physiology and phenology, also provide evidence for adaptation along elevation gradients. Together, these studies indicate that genetically based trait differentiation and adaptation to elevation are widespread in plants. We conclude that a better understanding of the mechanisms underlying adaptation, not only to elevation but also to environmental change, will require more studies combining the ecological and molecular approaches.     

Ethnobotanic garden design in the Ecuadorian Amazon

This study focuses on the documentation of traditional plant usage among Kichwa, the indigenous people from Canton Loreto, Ecuador. The relationship between people, plants and the natural environment is demonstrated in an ethnobotanical garden at the Capacity Building Centre of the town. The construction site for the ethnobotanical garden is a 1.5 ha secondary forest. The forest was analyzed with a local key informant and 150 different useful species were found. The plant species recorded are mainly used for medicinal purposes, followed by edible plants, and finally by food sources for animals. Open-ended interviews were conducted with the aim of identifying the most commonly used plant species among the Kichwas. The results showed that Ilex guayusa ranked most popular, followed by Myroxylon balsamum, Cedrela odorata, Banisteriopsis caapi, and Urera caracasana. Focus groups were held and the most important plant applications were evaluated. The collected data illustrated that Kichwas attach great importance to medicinal and ritual plants, followed by plants used for handcraft. Edible plants rank afterwards, followed by dye plants and plants used for hunting. The above findings serve as the backbone of the design for the ethnobotanical garden. The garden acts as a tool to preserve and promote the knowledge of plants, focusing mainly on medicinal plants. The growing areas for the plant species were determined according to their importance to the Kichwas. The concept of the ethnobotanical garden conveys the holistic picture drawn from the investigation on people and plants of the Kichwas.     

Geographic variations in phenotypic traits of the exotic herb Solidago altissima and abundance of recent established exotic herbivorous insects

Abstract

Many invasive plants increase aggressiveness after introduction. Since evolutionary forces such as herbivore pressure may change over different time scales, understanding the changes in biotic interactions in invasive plants through time can clarify the mechanism of their evolution in aggressiveness. In this study, we examined the geographic variation in phenotypic traits of Solidago altissima and the abundance of two exotic herbivorous insect species (the aphid, Uroleucon nigrotuberculatum and the lacebug, Corythucha marmorata), which are recently expanding their habitat on S. altissima populations over Japan. The two exotic insects were present at high density on S. altissima throughout their range. No differences in growth traits (plant height and number of leaves) were found among populations, and all plants examined appear to be exclusively hexaploid. Future studies on population genetics and common garden experiments are necessary to evaluate the potential evolutionary dynamics of the S. altissima after introduction.     

Partitioning the transplant site effect in reciprocal transplant experiments with Impatiens capensis and Impatiens pallida

Summary Modified reciprocal transplant experiments were conducted with the annual plant species Impatients capensis and I. pallida to partition the influence of environment on fitness into two components; that of (1) conspecific neighbours occupying each transplant site, and (2) the abiotic and biotic features of each site exclusive of the conspecific neighbours. In the within-species reciprocal transplant series, differences in survivorship and fruit production by cleistogamous flowers were attributable primarily to the effect of conspecific neighbours. In addition, plants surrounded by neighbours from the site of origin produced significantly more fruit from chasmogamous flowers compared with plants surrounded by neighbours from the alien site. In the between-species reciprocal transplant series, one transplant site was consistently associated with greater survivorship and fruit production regardless of the identity of neighbouring plants. The results suggest that different aspects of the environment in these reciprocal transplant experiments (conspecific neighbours, other species, physical factors) determine fitness in different situations.     

Variation in life-history traits among Urtica dioica populations with different history in parasitism by the holoparasitic plant Cuscuta europaea

Several studies demonstrate that natural enemies (e.g. parasites) have profound negative effects on the life-history traits of their hosts. If the host can compensate for the negative effects of parasitic infection by altering its life history, these modifications may partly form the basis of resistance or tolerance against parasites. Thus, parasites may be of considerable importance in shaping the evolution of life-history traits of their hosts. To examine if previous parasitism is associated with differences in life-history traits of the host, I conducted a common garden experiment with Urtica dioica plants originating from eight populations of which four were unparasitized, and four parasitized by the holoparasitic plant, Cuscuta europaea. A field survey indicated no differences between unparasitized and parasitized populations in, for example, the number of plant species and nutrient levels in the soil. Thus, it seems reasonable to assume that differences in life-history traits between the two population types in the common garden would reflect the effects of previous selection by the parasite. In the common garden, plants from parasitized populations started to flower later and allocated less biomass to asexual reproduction (measured as the production of stolons, i.e. clonal propagation) compared to plants from unparasitized populations. These results thus indicate that selection by the parasite may have favoured later onset of flowering, and may have selected against asexual reproduction.     

Effects of Competition and Life History Stage on the Expression of Local Adaptation in Two Native Bunchgrasses

Concerns about the use of genetically appropriate material in restoration often focus on questions of local adaptation. Many reciprocal transplant studies have demonstrated local adaptation in native plant species, but very few have examined how interspecific competition affects the expression of adaptive variation. Our study examined regional scales of adaptation between foothill and coastal populations of two California native bunchgrasses (Elymus glaucus and Nassella pulchra). By combining competitive manipulations with reciprocal transplants, we examined the importance of the vegetation at a site as a selective factor in the process of local adaptation. By monitoring survival and reproduction of reciprocally transplanted populations over the course of 3 years, we also studied the effect of life history stage on the expression of local adaptation. For most of the fitness components we measured, local adaptation was detected and interspecific competition consistently amplified its expression. Expression of local adaptation was especially apparent in the more inbreeding species E. glaucus and suggests that with weaker gene flow, selection may be more effective in creating ecotypes within this species. Local adaptation was detected at all life history stages but was most strongly expressed in traits associated with adult reproduction and the viability of seeds produced by the transplants. Taken together, our results indicate that the importance of local adaptation will become more apparent in the later stages of a restoration project as the plants at a site begin to reproduce and as they experience greater interspecific competition from the maturing vegetation at the site.     

Breeding system of the critically endangered Lakela’s Mint and influence of plant height on pollinators and seed output

Understanding reproductive systems of rare plants is critical for conservation efforts. Lakela's Mint, Dicerandra immaculata Lakela var. immaculata, is an endangered plant endemic to an approximately 4.8-km long area in Florida, USA. We used an experimental garden and three populations of Lakela's Mint to determine: (1) what is the breeding system (autonomous, asexual, self-fertile, cross-fertile) and are insects necessary for reproduction; (2) which native and nonnative insect species visit flowers and is the frequency of visits to a plant influenced by its height; (3) does the number of flowers visited within a plant by individual insects differ among native and nonnative insect species and due to plant height; and (4) is seed output influenced by plant height? Our results indicate that the breeding system of Lakela's Mint was facultative outcrossing. Insect-pollinated flowers produced more seeds than flowers that reproduced autonomously or asexually. The honey bee Apis mellifera L., a nonnative species, was the most frequent visitor to plants and visited more flowers within plants than native pollinators, but its behavior was not influenced by plant height. Native pollinators such as Bombus impatiens Cresson were attracted more frequently to shorter plants, but visited fewer flowers than on taller plants. Despite having fewer total and pollinated flowers, shorter plants had a higher output of intact seeds than taller plants, which could be due to differences in efficiency between native and nonnative pollinators or other factors. Our results add insight into factors influencing seed output and interactions between pollinators and rare plants.     

Seed source environment predicts response to water availability in Plantago patagonica

The intensity, duration, and severity of drought increasing across the American Southwest. Plant restoration efforts are often thwarted by drought-induced effects (i.e. drought-induced mortality or failure to reproduce). Careful selection of plant materials to match future environmental conditions could improve restoration success under climate change. Here, we focus on Plantago patagonica, a priority arid land restoration species in the southwestern US. Using experimental drought in a greenhouse and supplemental watering in a common garden experiment, we quantified how the seed-collection site environment influenced plant growth, performance, phenotypic plasticity, and evidence of local adaptation in 12 populations of P. patagonica. In the greenhouse, we found plants from hotter and drier environments had higher root:shoot ratios, while those from more variable precipitation regimes had greater total biomass in response to drought. Populations sourced from warmer environments exhibited sevenfold greater plasticity in root:shoot ratio compared to those from cooler environments, and there was strong evidence for local adaptation in phenology, as populations sourced from geographic locations nearer the common garden exhibited five times more flowering individuals than those sourced furthest from that garden. In the common garden, we found that plants sourced from wetter locations and those with more consistent precipitation regimes had lower mortality and higher specific leaf area, a proxy for growth, under unwatered ambient conditions. These results suggest plant performance of P. patagonica under reduced water availability is strongly shaped by the seed-collection site environment and may be a useful tool for improving restoration outcomes in a changing world.     

A test of the compensatory continuum: fertilization increases and below-ground competition decreases the grazing tolerance of tall wormseed mustard (Erysimum strictum)

Contrary to the general expectation, the compensatory continuum hypothesis proposes that grazing may not always affect plant performance adversely. Instead, the effects may vary from negative (undercompensation) to positive (overcompensation), depending on the local availability of resources and the intensity of competition experienced by individual plants. We tested this hypothesis in a common garden experiment by growing tall wormseed mustard, Erysimum strictum, under a factorial design involving simulated grazing (0, 10, or 50% of the main stem clipped), supplemental fertilization and below-ground competition. The results supported the hypothesis. On an average, fertilization increased and competition decreased plant performance. Overcompensation was only observed among the fertilized plants growing free of competition. Simulated grazing increased seed yield 1.6 (10% clipping) and 1.4 times (50% clipping) as compared to unclipped plants when the plants were grown with fertilization and without competition. In contrast, clipping did not significantly increase seed yield in the plants grown without fertilization and/or with competition. The breakage of apical dominance provides a proximate mechanism of these regrowth responses. This is consistent with the fact that most plants (85%) had an unbranched shoot architecture in our study population. However, it is not clear why E. strictum has a relatively unbranched architecture in natural populations. We briefly discuss the alternative ecological factors – competition for light, adaptation to herbivory and optimal timing of flowering as a bet-hedging strategy in monocarpic plants – which might maintain unbranched architecture in this species. This revised version was published online in July 2006 with corrections to the Cover Date.