Density and the commitment of apical meristems to clonal growth and reproduction in Hieracium pilosella

Summary We examined responses to population density in the commitment of apical meristems to reproduction and clonal growth in a rosette-forming, stoloniferous herb (Hieracium pilosella). Despite close physiological coupling between the evocation of the terminal inflorescence bud and the development of one or more axillary buds into stolons, the allocation of meristems was extremely plastic.Genets at the higher sowing densities showed density-dependent mortality consistent with self-thinning along a-3/2 trajectory. The probability of inflorescence evocation and associated stolon development was negatively dependent on surviving density. The proportinal distribution of primary stolons amongst genets became strikingly more unequal (expressed as the Gini coefficient) with increasing density. Clonal growth was resolved into the number of primary stolons per stoloniferous genet and the extent of stolon branching (i.e. number of apices per primary stolon); both showed strongly negative density-dependence. Reproduction, expressed as the mean number of flowering capitula per stoloniferous genet, declined 15-fold with increasing density; although theoretically expected to be unity, greater values resulted from capitulum production by attached secondary rosettes and lower values reflected the increasing abortion rate of inflorescence buds with increasing density.Both the total number of apices produced per unit area and the corresponding number of reproductive apices were maximal at intermediate surviving densities (700–1,000 m^-2). The balance between reproductive and clonal growth may be expressed as the probability of an apical meristem producing a capitulum, that also peaked sharply at intermediate density. This finding does not conform with linear models that predict a shift from vegetative growth to sexual reproduction with increasing population density.     

Habitat protection,cattle grazing and density‐dependent reproduction in a desert tree

Anthropogenic activities usually trigger changes in the population density of plants. Thus, land management practices can influence density‐dependent demographic parameters and species interactions. We investigated plant‐pollinator interactions and reproduction in Prosopis flexuosa, the largest tree species in the Central Monte desert of Argentina, an important economic and cultural resource for humans and a functionally prominent species. We hypothesized that reproductive output of P. flexuosa would be limited at low densities, and that exclusion of catle grazing would enhance population density and consequently interaction frequency with pollinators and reproductive success. The study was conducted in and around Ñacuñán Biosphere Reserve (Mendoza, Argentina), where cattle grazing has been excluded for over 35 years. Working in five pairs of protected and cattle grazed 1‐ha plots, we recorded density of adult trees, pollinator visitation frequency to inflorescences and seeds per inflorescence in focal trees. Adult tree density was higher in protected plots than in cattle grazed plots. Density of reproductive trees was positively correlated with seed production, suggesting positive density dependence for reproduction (Allee effect). Pollinator visitation to inflorescences and seed production was higher in protected plots compared with plots under cattle grazing. Suppression of anthropogenic degradation has resulted in higher adult tree density in protected plots, indirectly higher pollinator visitation to inflorescences and higher reproductive success of trees. Increased frequency of plant‐pollinator interactions and tree reproduction suggest success of management practices aimed at protecting P. flexuosa woodlands.     

Resource sharing among ramets in the clonal herb,Fragaria chiloensis

Summary The herbaceous perennial, Fragaria chiloensis, reproduces vegetatively on coastal sand dunes in California by growth of stolons that bear rosettes. Movement of water and photosynthates through stolons integrates water and carbon metabolism of rosettes both before and after they root. New, unrooted rosettes import sufficient water and nitrogen to maintain levels near those of established rosettes; yet support of an unrooted rosette did not decrease growth of a connected, rooted sibling given abundant light, water, and soil nutrients. Under such conditions strings of unrooted rosettes with the associated stolon appeared self-sufficient for carbon; shade and drought induced import of photosynthates. New rosettes produced and maintained a limited root mass upon contact with dry sand, which could increase probability of establishment. Rooting did not induce senescence of stolons. Connection between two established rosettes prevented death by drought and shade, even when neither rosette could have survived singly. Results suggest that physiological integration of connected rosettes may increase total growth of clones of F. chiloensis through sharing of resources among ramets, especially when resource availability is changeable or patchy.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

Does fluctuating resource availability increase invasibility? Evidence from field experiments in New Zealand short tussock grassland

The theory of fluctuating resource availability proposes that the susceptibility of a plant community to invasion by new species (i.e., invasibility) depends upon conditions of intermittent resource enrichment coinciding with the presence of invading propagules. We compared the response of a rapidly invading forb (Hieracium pilosella L.) between different experimental treatments in a short tussock grassland in New Zealand, over 6–12 years, to determine whether the theory explains differences in invasibility. The theory predicts that environments subject to periodic resource enrichment will be more invasible than those with more stable resource-supply rates. In our study, H. pilosella did not increase more rapidly in treatments subject to periodic resource pulses (fertiliser and water) than in those with more stable resource supplies. Also contrary to the predictions of the theory, the rate of invasion of H. pilosella did not increase following an increase in the rate of supply of water or nutrient resources, or following treatments that temporarily reduced resource uptake in the community, including grazing. H. pilosella did not increase immediately following abrupt increases in water and nutrient supply and removal of the dominant grass species with herbicide, as predicted by the theory, although temporary increases in resident exotic guilds indicated that the intensity of competition for resources was reduced. Neither H. pilosella nor resident exotic guilds showed increased cover growth rates following resumed grazing. The rate of invasion by H. pilosella was not correlated with species richness, a result consistent with one of the predictions of the theory. Therefore, short-lived events that temporarily reduced or suspended competition did not appear to determine the invasion success of this particular species in this region. In New Zealands perennial short tussock grasslands, the characteristics of the resident plant community may be more critical than resource fluctuations in determining invasion success of H. pilosella. Invasion of H. pilosella may be most successfully controlled here by promoting a successional physiognomic shift to a taller, shrub-and-tussock-dominated canopy that competitively excludes low-growing forbs.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

Germination and population structure of spear thistle Cirsium vulgare in relation to experimentally controlled sheep grazing

Summary The size and number of rosettes of Cirsium vulgare were censused in a 4 ha sheep grazing trial on lowland calcareous grassland in paddocks receiving controlled winter, spring and summer grazing treatments. Spring grazing significantly increased thistle rosette numbers, and there were always fewer rosettes in size classes 250mm–300 mm under the heavier grazing treatments. The emergence of seeds of Cirsium vulgare sown by hand into each of the grazing treatments was monitored and showed a positive effect of spring grazing. When the percent emergence of sown seeds was used as a covariate in the analysis of rosette numbers in the experimental paddocks, it was found to account for 77% of the variance in thistle numbers. It was concluded that the main effect of grazing on thistle rosette numbers is an indirect one exercised through the effects of grazing on germination conditions. Seeds of Cirsium vulgare were sown into artificial gaps at another grassland site, to determine the effect of gap-size upon seedling emergence. Germination was poor, but significantly more seedlings emerged in gaps 10–20 cm diameter, than in 5 cm gaps or in control plots without a gap. A computer simulation model was used to explore the relationship between gap density and thistle population dynamics. A threshold density of gaps was found to exist, below which thistle populations went extinct, and above which the thistle population grew geometrically. The degree of aggregation of dispersed seeds did not alter the threshold gap-density for plant extinction, but did affect the rate of increase of the thistle population when the threshold gap-density was exceeded. It is concluded that strategies for controlling Cirsium vulgare populations by grazing manangement will be most effective if aimed at reducing suitable sites for establishment in spring. The success of such attempts will depend upon the soil fertility of the site, and sudden outbreaks of Cirsium vulgare infestation can be expected if a sward gradually deteriorates through over-grazing.     

A comparison of native and invasive populations of three clonal plant species in Germany and New Zealand

Aim Our aim was to test for changes in growth patterns of three clonally growing plant species (Achillea millefolium, Hieracium pilosella and Hypericum perforatum) between native and invaded regions. We addressed the hypotheses that with differing important life‐history traits, invasive populations perform better than native populations, and that this expected better performance is linked to weakened trade‐offs between individual growth and sexual and clonal reproduction. Location Germany and New Zealand. Methods We conducted field surveys for the three above‐mentioned species in both native German and invasive New Zealand populations, and collected data at both population and individual levels. Results At the population level, the proportion of flowering plants, population size and population density were all higher in invasive populations. Similarly, at the individual level, the number of stolons per plant, stolon–biomass ratio and population crowdedness (local plant density in a specified area around a target plant) were significantly higher in New Zealand. Plant height did not differ between countries, and plant biomass was lower in New Zealand than in Germany for Achillea millefolium and Hypericum perforatum. These two species showed significant trade‐offs between individual growth and sexual and clonal reproduction. Achillea millefolium exhibited a weakened trade‐off in its invaded range, where the same proportion of flowering plants was sustained at much higher levels of population crowdedness than in its native range. Main conclusions The apparent invasion success of the three study species is generally due to better overall performance in their respective invaded ranges. In respect of both Achillea millefolium and Hypericum perforatum, this is driven primarily by increased vegetative reproduction. In contrast, Hieracium pilosella seems to benefit more from increased sexual reproduction in its invaded range. Shifts in trade‐offs as a general trend seem to be of minor importance.     

Factors limiting rosette recruitment in scarlet gilia, Ipomopsis aggregata: seed and disturbance limitation

We performed a factorial combination of seed additions, surface soil disturbance, and protection from potential seed predation in experimental plots to elucidate the relative importance of each factor in the recruitment of scarlet gilia seedlings. We detected a significant interaction between seed addition and surface soil disturbance on seedling recruitment over 2 years of establishment. Plots that received both supplemental seed addition and disturbance established 10.5 times as many rosettes as control plots. We did not detect main or interactive effects of caging on seedling establishment. We explored the importance of density-dependent recruitment by investigating the number of rosettes per experimental plot through time. In addition, we used planned contrasts to compare the absolute and percent changes in control and treatment plot rosette densities between the study years. We found significant downward concave curvature to the seedling recruitment curve, suggesting negative density-dependent recruitment across the experimental range of rosette densities. We found a significant difference between the control plots and the seed addition-disturbance plots in terms of absolute changes in plot density from 1997 to 1998. Greater net mortality occurred in plots that received both seed addition and disturbance. However, we found no differences among treatments in the percent change in rosette density across the study years. Our study demonstrates that seedling recruitment in natural plant populations may be limited by the interaction of seed and microsite availability, and that seedling recruitment in scarlet gilia may be negatively density dependent. Our results suggest that the fecundity effects of particular plant-animal interactions (e.g., pollination, herbivory) may affect scarlet gilia population dynamics, particularly under conditions of high soil disturbance. Received: 11 June 1999 / Accepted: 24 November 1999     

Combined effects of folivory and neighbor plants on <Emphasis Type="Italic">Cirsium altissimum</Emphasis> (tall thistle) rosette performance

Predicting how herbivory and neighbor plant interactions combine to affect host plants is critical to explaining variation in herbivores’ impact on plant population dynamics. In a field experiment, we asked whether the combined effects of neighbor plants and folivores upon performance of tall thistle (Cirsium altissimum), a monocarpic perennial, can be predicted as the product of their individual effects (i.e., effects of neighbor plants and folivores act independently in suppressing tall thistle performance). Alternately, the combined effects of neighbor plants and folivores might be greater, indicating a synergistic interaction, or less, indicating an antagonistic interaction, than the product of their individual effects. Our experiment involved a neighbor plant clipping treatment and a folivory reduction treatment in a factorial design with manipulations applied to naturally occurring tall thistle rosettes in restored tallgrass prairie. Clipping neighbors at the soil surface within 40 cm of tall thistle rosettes increased light availability to rosettes, rosette growth, and the transition rate of 2007 rosettes to reproductive adults in 2008. Folivores’ and neighbor plants’ effects acted independently upon rosette growth. By contrast, folivory reduced the rate at which 2007 rosettes transitioned to reproductive adults in 2008 only where neighbor plants were unclipped, indicating a possible synergistic interaction of neighbor plants and folivores in suppressing tall thistle performance. Our results suggest that (1) promoting neighbor plant aboveground biomass should suppress rosette-forming weeds, and (2) folivory, which reduces light acquisition by rosettes, may generate synergistic herbivory × neighbor plant interaction effects on rosettes in grasslands, where light often limits rosettes.     

Population recovery of the sea urchin Lytechinus variegatus in a seagrass flat (Araruama Lagoon,Brazil): the role of recruitment in a disturbed environment

Summary

The size, structure, and density of a population of Lytechinus variegatus were recorded from a shallow seagrass bed (Halodule wrightii) at 40-day intervals from April 1994 to April 1995. A local mass mortality was observed in May 1994. The density declined from 0.57±0.10 to 0.06±0.05 ind.m^?2, mean size declined from 37.6±0.49 to 16.2±2.14 mm and maximum size from 75 to 40 mm. Ten months after mortality, the maximum and mean sizes had returned to initialvalues, but density had not (0.10±0.03 ind.m^?2). A second, smaller mass mortality occurred in April 1995. High values of gonad index were recorded before mortality (9.40±5.97) and after population recovery (11.96±5.74). Recruitment was continuous throughout the year, but reached a peak from August to October. This can he related to the patterns of upwelling in the region. Mass mortality is probably an annual event in this population that can be correlated with the occurrence of low tides during daytime in the fell. The population can recover during the summer when low tides occur during the night. The massive input of recruits prevents extinction of the population during the season of high mortality. Population recovery depends on the magnitude of recruitment.     

Growth and Development in Arabidopsis thaliana through an Entire Life Cycle under Simulated Microgravity Conditions on a Clinostat

Arabidopsis thaliana ecotype Columbia and Landsberg erecta were studied. Horizontal clinorotation affected little germination of seeds, growth and development of rosette leaves and roots during early vegetative growth stage, and the onset of the bolting of inflorescence axis and flower formation in reproductive growth stage, although it suppressed elongation of inflorescence axes. The clinorotation substantially reduced the numbers of siliques and seeds in Landsberg erecta, and completely inhibited seed production in Columbia. Seeds produced in Landsberg erecta on the clinostat were capable of germinating and developing rosette leaves normally on the ground. On the other hand, growth of pin-formed mutant (pin/pin) of Arabidopsis ecotype Enkheim, which has a unique structure of inflorescence axis with no flower and extremely low levels of auxin polar transport activity, was inhibited and the seedlings frequently died during vegetative stage on the clinostat. Seed formation and inflorescence growth of the seedlings with normal shape (pin/+ or +/+) were also suppressed on the clinostat. These results suggest that the growth and development of Arabidopsis, especially in reproductive growth stage, is suppressed under simulated microgravity conditions on a clinostat. To complete the life cycle probably seems to be quite difficult, although it is possible in some ecotypes. Received 18 June 1999/ Accepted in revised form 27 August 1999     

From horticultural plantings into wild populations: movement of pollen and genes in <Emphasis Type="Italic">Lobelia cardinalis</Emphasis>

The Southern Grassland biome in Brazil has long been neglected by the government and by the scientific community as well. Lack of studies about the effects of disturbance (mostly fire and grazing) on plant community and populations are the main hindrances for the future management plans, which complicates the conservation of this rich and unique biome. In this study, the population biology of a rosette species, Eryngium horridum (Apiaceae), has been investigated, in order to compare the effects of fire, grazing and abandonment. E. horridum is a very common species and many farmers consider it a nuisance. Two different areas were chosen for the study purposes: Morro Santana and the Research Center Pró-Mata. Areas under different types (fire and grazing) and disturbance histories were compared. Populations were analysed in plots of 25 m², divided in age-state classes (seedlings, young, adult and reproductive individuals). Both morphological and phenological traits have been investigated. Fire influenced populations in a direct way, whilst grazing showed an indirect effect. Seedlings were found only in areas excluded from disturbance for 3 years on Morro Santana. Due to this, seedling establishment might be more related to environmental conditions than to disturbance regimes. Young individuals were mainly found in recently burned areas, while adult individuals were dominant in areas excluded from disturbance. Additionally, individuals from E. horridum showed a great capacity of resprouting after plant damage. Reproductive individuals also showed the capacity of producing new rosettes after the death of the inflorescence axis. This strategy is probably of great importance for the survival of populations in areas excluded from disturbance. Therefore, vegetative reproduction seemed to be the most important strategy for the maintenance and survival of its populations, allowing longer population persistence over time, even in the absence of disturbance.     

The cost of reproduction in Senecio keniodendron,a giant rosette species of Mt. Kenya

Summary A three year study of Senecio keniodendron (Compositae), a giant rosette species of the alpine zone of Mt. Kenya, demonstrated that individuals which reproduce are more likely to die, and less likely to reproduce in the future if they do survive, than are vegetative individuals of the same size. However, if an individual reproduces, survives and reproduces again, then it produces more seeds during the second reproductive episode than does a plant of the same height reproducing for the first time, because reproduction is followed by production of lateral rosettes, increasing the number of potentially-reproductive rosettes per plant.Slow-growing rosettes are less likely to reproduce than fast-growing rosettes. For rosettes which do reproduce, rosette size and rate of leaf production, measured before reproduction begins, are good predictors of fecundity.     

Density-dependent variation in reproductive success in a terrestrial orchid

 Within population variation in plant density can affect reproductive success and breeding systems. We examined such effects in Listera cordata R. Br., a north temperate terrestrial orchid, in a subalpine population in Colorado, USA. Hand pollinations showed that L. cordata was self-compatible as all pollinations produced fruits. Cross-pollinations differed significantly from self-pollinations, and had a higher number of seeds per capsule and higher % of fertilized ovules. Together they could compensate for the transmission advantage of self-pollinations. Average number of flowers per inflorescence was 4.2–4.4 and average fruit set was 20.7–23.7%. By establishing 36 plots with varying plant densities (1, 4, 8 and 16) and emasculating all flowers in target plants of a set of 20 plots, we estimated density effects on fruit and seed production. Plant density had no effect on the amount of reproduction except in reduced seed production of highly dispersed plants. Our results suggest that inbreeding depression may be an important selective factor in L. cordata. Further studies are needed to define its role in the evolution of mating systems in this species. Received February 21, 2000 Accepted December 26, 2000     

Turfgrass (Cynodon dactylon L.) sod production on sandy soils: II. Effects of irrigation and fertiliser regimes on N leaching

The effects of irrigation and fertiliser regimes on N leaching from the production of couch grass (Cynodon dactylon L.) sod, on a free-draining sandy soil, were evaluated in a 22-month field study. The experimental design used a randomised-block, split-plot design with three replicates. Main plots consisted of two irrigation treatments: 70 and 140% daily replacement of pan evaporation; four subplot fertiliser types (water-soluble (predominately NH₄NO₃), control-release, pelletised poultry manure and pelletised biosolids); and three N application rates (100, 200 and 300 kg N ha⁻¹ per crop). Nitrogen leaching was assessed by measuring the leachate volumes and concentrations of N species leached from soil lysimeters (250 mm in diameter by 950 mm in length) installed in 10 m² turfgrass plots. Nitrogen leaching ranged from 33 to 167 kg N ha⁻¹ over 22 months, depending upon the irrigation and fertiliser treatment. Irrigation treatment affected N leaching more than fertiliser treatment, and increasing the irrigation from 70 to 140% replacement of daily pan evaporation increased N leaching for all fertiliser types, and by up to four times. Forty six to 76% of losses occurred from the high irrigation treatments during the first 16 weeks after the turfgrass was planted as rhizomes. By contrast, N leaching did not appear to increase following harvest of sod. At the high irrigation treatment, N leaching was greater for the pelletised biosolids than the control-release; while at the low irrigation treatment, N leaching did not vary between fertiliser types. A significant proportion of the N leached was in the organic form. Therefore, we recommend total N and mineral N be measured when assessing N leaching from turfgrass. Nitrogen leaching from turfgrass production is low from all fertiliser types when the irrigation matches turfgrass water use and N is applied at a rate and frequency that approximates turfgrass requirements. Section Editor: P. J. Randall     

Habitat characteristics,stage structure and reproduction of colline and montane populations of the threatened species <Emphasis Type="Italic">Arnica montana</Emphasis>

Arnica montana, a characteristic plant of nutrient-poor grasslands, has strongly declined during the last decades, particularly at lower altitudes. To gain insights into the underlying reasons for this decline, we recorded differences in size, stage structure and reproductive traits between 21 colline populations of A. montana in the Ardennes-Eifel region (280–600 m a.s.l.) and 10 montane populations in the Vosges Mountains (1,200 m a.s.l.). Community composition of the Violion caninae vegetation at colline sites and Nardion strictae vegetation at montane sites indicated that temperature and moisture were the main abiotic factors differentiating between low and high altitudinal relevées. The proportion of flowering rosettes decreased with altitude, indicating a shift from sexual reproduction to clonal growth. In contrast to expectation, Ellenberg values for nutrient availability as well as the proportion of young rosettes and population size did not differ between colline and montane populations. However, population size decreased with nutrient availability, indicating eutrophication as the reason for population decline. In small populations, plant density, the proportion of flowering rosettes, the number of flowerheads per rosette and the number of seeds per flowerhead were lower, indicating less suitable conditions. Thus, preventing further eutrophication in both colline and montane populations will be crucial for the conservation of the species. Reduced reproduction may not be important for the population dynamics of this clonal plant in the short term, but could affect genetic diversity and survival in the long term.     

Dry soil determines the pseudoviviparous reproduction in Comanthera nivea (Eriocaulaceae) in the Campos Rupestres in south-eastern Brazil

Pseudovivipary is an asexual reproduction strategy correlated with extreme environments. Comanthera nivea occurs in habitats with low water retention. This study investigated the effect of soil moisture availability on C. nivea pseudoviviparous reproduction and flowering. We established four permanent plots (1 × 1 m), two in a shaded area, in which we marked 15 rosettes in each plot, and two in a full-sun area, in which we marked 10 rosettes in each plot. We made monthly observations from August 2016 to January 2017, to quantify the number of flower heads and of pseudoviviparous ramets per rosette, and measured soil moisture. Our results showed a high correlation between soil moisture and flowering. Flowering in C. nivea occurs in periods of increased soil moisture, whereas pseudoviviparous reproduction occurs in the driest period. The advantages of a pseudoviviparous canopy-forming strategy in habitats with scarcity of water in the soil are discussed.     

Life history and population dynamics of Atriplex triangularis

The dynamics of a population of the stoloniferous Potentilla anserina was studied during four years. The mortality of established ramets followed a log-linear pattern superimposed on a seasonal fluctuation with an increased mortality during the period of vegetative spread. The major external mortality factors were unfavourable weather conditions such as drought and freezing. The turnover rate of the population was slow (31.6 yr). Vegetative spread dominated the recruitment to the population. Seedlings occurred but were exposed to a much higher mortality than vegetative daughter ramets. The juvenile period of seedlings is at least 4 yr in the field. This contrast to vegetative daughter ramets which are potentially flowering and stoloniferous the year after they are established, even if age affects both reproduction and vegetative spread. The density of most of the population is below the level where intraspecific regulative effects operate. However, in very dense stands both stolon and flower production were negatively density dependent.Nomenclature follows Lid (1974), Norsk og Svensk Flora, 2nd ed. Oslo.I am indebted to Lenn Jerling, Peter Torstensson, Anders Telenius and Gudrún Jónsdóttir for their help, and inspiring criticism throughout this study.     

Effects of local density and flower colour polymorphism on pollination and reproduction in the rewardless orchid Dactylorhiza sambucina (L.) Soò

The rewardless orchid Dactylorhiza sambucina shows a stable flower colour polymorphism, with both yellow- and red-flowered morphs growing sympatrically. Pollination biology and breeding system were investigated to examine the effects of density of plants, colour polymorphism, inflorescence dimension, and flower position within inflorescence on male and female reproductive success in three natural populations of D. sambucina. There were significant differences among sites in the number of pollinia removed and in fruit set per inflorescence. Number of removed pollinia and capsule production in D. sambucina were independent from flower and inflorescence size or flower position. As a whole, the red morphs showed the highest number of capsules produced, while the yellow morphs had the greatest male success. The relative male and female reproductive success were independent from plant density but were significantly correlated with the yellow morph frequency at the population level. Overall, our findings show that the contribution to the total reproductive success deriving from the two colour morphs does not conform with the predictions of negative frequency-dependent selection.