Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.     

Microsatellite loci in eelgrass Zostera marina reveal marked polymorphism within and among populations

Using an enriched genomic library, we developed seven (CT)n/(GA)n microsatellite loci for eelgrass Zostera marina L. Enrichment is described and highly recommended for genomes in which microsatellites are rare, such as in many plants. A test for polymorphism was performed on individuals from three geographically separated populations (N = 15/population) and revealed considerable genetic variation. The number of alleles per locus varied between five and 11 and the observed heterozygosities for single loci ranged from 0.16 to 0.81 within populations. Mean allele lengths were markedly different among populations, indicating that the identified loci will be useful in studying population structure in Z. marina. As the frequency of the most abundant multilocus genotype within populations was always < 1%, these loci have sufficient resolving power to address clone size in predominantly vegetatively reproducing populations.     

Effect of flowering on vegetative growth and further reproduction in

Flowering intensity and plant size were monitored in 155 Festuca novae-zelandiae individuals over four years to determine if trade-offs exist between inflorescence production and vegetative growth, and between inflorescence production in different years. Less than half of the population flowered in any one year, 36% of individuals did not flower at all, and only 17% flowered in all four years of the study. Mean number of inflorescences per individual per year varied from 1.54 to 5.53 (maximum = 85). No trade-offs were detected between flowering frequency and intensity; individuals that flowered more frequently also produced more inflorescences in each flowering episode. No trade-off was detected between current and future reproduction, rather flowering intensity was positively correlated between years. Growth, as measured by diameter increment, was positively related to flowering frequency and flowering intensity, both across all individuals studied and within 1m x 1m plots. The presence of a positive relationship between growth and reproduction within plots argues against meso-scale variability in environment factors being the cause of the results from analyses involving all individuals. Clearly reproduction in F. novae-zelandiae does not incur a marked cost in growth or future reproduction. The assumptions underlying theoretical expectations of such trade- offs may not be valid for long-lived clonal plants such as F. novae-zelandiae.     

Population ecology in natural<Emphasis Type="Italic">Syneilesis palmata</Emphasis> stands: I. relationship between flowering and plant size

In this study, the relationship was explored between flowering potential and plant size in a population ofSyneilesis palmata (THUNB.) MAX., Usannamul. For this perennial shade-tolerant herb, the flowering rate, plant size, and changes that occurred in the next year after flowering were monitored in two different types of natural stands: moderately shaded (MS; for 11 years) and severely shaded (SS; 7 years). The correlation coefficients (CC) between inflorescence number and other factors, i.e., widths of the first leaf, second leaf, or total leaf breadth, were significant at the 1% level in both stands. In particular, the CC was >0.9 between the number of inflorescences and total leaf breadth. Although the mean flowering rates averaged 6.8% in MS and 8.6% in SS, those values varied widely, ranging from 0 to 53% among years. These rates were not related to petiole length but did increase in proportion to the width of the leaf blade. Flowering began when plants moved beyond the medium size class; the rate in the MS environment was 100% for the largest-sized plants. For plants in the SS, however, the highest rate was 35%. Over the 11 years, 62.8% of all plants in the MS never flowered, while another 30.8% bloomed only once. In comparison, plants observed for 7 years in the SS showed a non-flowering rate of 78.1% and a one-time-only rate of 19.2%. In the next year after flowering, 57.6% of the MS plants exhibited reduced growth while 25.8% of those individuals each split into two plants. In the SS, 37.8% had decreased development the following year, and a mortality rate of 37.8%.     

Spatiotemporal variation in predispersal seed predation intensity

The effect of predispersal seed predation by Bruchus atomarius (Bruchidae, Coleoptera) on individual performance and population dynamics of the perennial forest herb, Lathyrus vernus (Leguminosae), was investigated in 11 permanent plots over 4 years. Seed predation and parameters describing intra-specific neighbour distance, plant size, inflorescence size, flowering phenology and current and previous herbivore damage were measured on all plants. In addition, demographic information from all plots was analysed using transition matrix population models in order to estimate the influence of seed predation on population growth rates. Predispersal seed predation rates differed significantly among years. Plot averages ranged from 0 to 83.7%. However, most of the variation occurred among individuals. Within individuals there was no consistency in predation rates among years. Exposure to herbivory, plant size and flowering phenology did not affect predation rates but individuals with larger inflorescences suffered from significantly higher predation. Seed predation in L. vernus was not influenced by neighbour distances of individual plants but it was positively correlated with the average density of seeds within plots, suggesting that seed predation is density dependent at the patch level. The reduction in population growth rate due to seed predation ranged from 0 to 7.6%. The sensitivity of population growth rate to reductions in seed production varied considerably among years and plots. This variation was mainly due to differences in the reproductive value of seeds and seedlings. The intensity of seed predation over the range found was not correlated with changes in population growth rate. The results of this study suggest that the influence of external factors, like seed predation, on population growth rate largely depends on the demographic transition rates in the investigated population.     

高山垫状植物团状福禄草开花面积与方位随海拔的变化及其适应性

作为高山生态系统中的奠基种(foundation species), 垫状植物自身种群的繁殖与扩张, 对高山生态系统功能稳定性起着关键作用。但是, 垫状植物如何在极端环境条件下实现资源的有效利用与分配, 达到繁殖最优化, 至今鲜为人知。该研究在滇西北白马雪山沿海拔梯度选择具有不同坡度及坡向的5个团状福禄草(Arenaria polytrichoides)种群, 调查、比较种群内、种群间以及具有不同性系统的植株个体之间的开花面积比、开花方位, 并分析不同生态因子对其开花特性的影响。结果表明: 随着海拔的升高, 团状福禄草个体变小, 其分配到开花的资源比例总体上随海拔上升呈现下降的趋势, 说明团状福禄草的繁殖分配受到由海拔所引起的生态因子的调控。但是, 部分低海拔种群内植物个体的繁殖分配显著低于部分高海拔种群, 说明海拔并非控制植物繁殖分配的唯一因素。此外, 植株开花总面积随植株个体增大而增加, 但开花面积比却随个体增大而变小, 说明植株分配到开花的资源增长速率可能低于植株个体的增长速率。在性别差异方面, 两性植株对开花的资源分配比例要显著高于雌性植株, 但是, 其差异程度受到海拔因素的影响。最后, 在同一种群内, 团状福禄草在冠层表面不同方位上的开花面积比存在显著差异性, 这种差异性在不同种群之间又具有不同的表现形式。     

Is increased male flower production a strategy for avoidance of predispersal seed predation in andromonoecious plants?

Floral gender in angiosperms often varies within and among populations. We conducted a field survey to test how predispersal seed predation affects sex allocation in an andromonoecious alpine herb Peucedanum multivittatum. We compared plant size, male and perfect flower production, fruit set, and seed predation rate over three years among nine populations inhabiting diverse snowmelt conditions in alpine meadows. Flowering period of individual populations varied from mid‐July to late August reflecting the snowmelt time. Although perfect flower and fruit productions increased with plant size, size dependency of male flower production was less clear. The number of male flowers was larger in the early‐flowering populations, while the number of perfect flowers increased in the late‐flowering populations. Thus, male‐biased sex allocation was common in the early‐flowering populations. Fruit‐set rates varied among populations and between years, irrespective of flowering period. Fruit‐set success of individual plants increased with perfect flower number, but independent of male flower number. Seed predation by lepidopteran larvae was intense in the early‐flowering populations, whereas predation damage was absent in the late‐flowering populations, reflecting the extent of phenological matching between flowering time of host plants and oviposition period of predator moths. Seed predation rate was independent of male and perfect flower numbers of individual plants. Thus, seed predation is a stochastic event in each population. There was a clear correlation between the proportion of male flowers and the intensity of seed predation among populations. These results suggest that male‐biased sex allocation could be a strategy to reduce seed predation damage but maintain the effort as a pollen donor under intensive seed predation.     

Altitudinal variation in flowering area and position and their ecological significances of an alpine cushion Arenaria polytrichoides,a gynodioecious herb

Aims As foundation species in the alpine ecosystems, the reproduction and recruitment of alpine cushion plants are very important for sustaining the alpine ecosystem functions. However, it still remains unclear that how cushion plants effectively allocate resources to optimize reproductive fitness.Methods Here we selected five populations of a gynodioecious herb Arenaria polytrichoides with different exposures and slopes along an altitudinal gradient on the Baima snow mountain in northwest Yunnan, southwest China, to investigate and compare flowering area and positions, within and among populations and between female and hermaphroditic morphs. By doing so, we further discuss how the environmental stresses affect the cushion’s flowering attributes thus the population-level reproduction.Important findings The results showed that, individual plant size and resources allocated to flowering (flowering area %) both decreased with increasing elevation, indicating that the reproductive allocation strategy was significantly affected by elevation. However, a population at lower elevation showed lower reproductive investment than higher populations, suggesting that elevation was not the only factor affecting the cushion’s reproductive allocation. In addition, absolute flowering area increased with increasing individual size, but the flowering area ratio decreased, indicating that the increases in reproductive allocation are fewer than that in vegetative allocation. Hermaphroditic individuals invested more resources to flowering than females did, but again, such effect was affected by elevation. Moreover, within a single population, the flowering areas were significantly different among the four directions (east, south, west and north) within one single individual canopy, but such differences varied in different populations.     

Density-dependent seed set in the Haleakala silversword: evidence for an Allee effect

Plant species may be subject to Allee effects if individuals experience a reduction in pollination services when populations are small or sparse. I examined temporal variation in reproductive success of the monocarpic Haleakala silversword (Argyroxiphium sandwicense subsp. macrocephalum) over five years, to determine if plants flowering out of synchrony with most of the population (i.e., in low flowering years) exhibited lower percent seed set than synchronously-flowering plants (i.e., those flowering in high flowering years). Through two pollination experiments conducted over multiple years, I also measured pollen limitation and self-incompatibility in this species. The number of flowering plants varied greatly among years, as did reproductive success. Percent seed set was significantly correlated with the number of plants flowering annually, such that plants flowering in high flowering years (1997 and 2001) exhibited significantly higher percent seed set than did plants flowering in low flowering years (1998-2000). In the 3-year pollen limitation study, plants flowering asynchronously were pollen-limited, whereas plants flowering synchronously were not. This species is strongly self-incompatible. Results of this study demonstrate that the Haleakala silversword experiences reduced reproductive success in low flowering years, and suggest that this Allee effect is pollinator-mediated. Allee effects in plants are an understudied yet potentially important force with implications for the population dynamics and conservation of rare species.     

INDIVIDUAL FLOWERING TIME IN A GOLDENROD (SOLIDAGO CANADENSIS): FIELD EXPERIMENT SHOWS GENOTYPE MORE IMPORTANT THAN ENVIRONMENT

Individual plants of an old field population of the clonal perennial goldenrod, Solidago canadensis L. are consistent in rank order of flowering time over years. In order to analyze whether this consistency is due more to environmental variations among microsites of individual clones or due to genetic variation among clones, similar-sized sections of rhizomes were transplanted to an experimental garden in a randomized block design with five replicates per clone. Three years later we examined the transplants and the original plants for their flowering phenology. In the experimental garden there was an unexpected soil moisture gradient across blocks, with associated gradients in height of goldenrods and height of surrounding vegetation. No gradient was significantly correlated with the flowering time of the transplants. That is, the time of flowering of transplants was consistent among replicates of individual clones across the soil moisture gradient and regardless of the size of the transplants themselves, or the height of surrounding vegetation. Further, the flowering times of the transplants were significantly positively correlated with the flowering times of the original plants in the same year. We conclude that the differences in flowering time among clones within a goldenrod population is largely determined by genetic variation, rather than by environmental factors.     

Within and Between Plant Variation in Seed Number, Seed Mass and Germinability of Primula elatior: Effect of Population Size

Abstract: In this study, natural variation of seed mass and seed number between and within individual plants of P. elatior, a common forest herb on rich soils in Flanders, was investigated for both small (< 50 flowering individuals) and large (> 1000 flowering individuals) populations. The relationship between seed number and seed mass at both the between and within population level was also quantified. We also investigated how population size affected germinability of seeds through the effects of seed mass. Seed number varied by a factor of twenty, whereas seed mass spanned, on average, a 3.5 range. In both cases variation was highest among populations, indicating that population size has important effects on seed number and seed mass. Next, seed number-seed mass tradeoffs were significantly influenced by population size, with small populations showing strong negative relationships whereas in large populations seed mass was only marginally influenced by seed number. As to germination, larger seeds showed larger germination percentages only for the smallest population. In all other populations, seed mass had little effect on germination.     

Population biology of the rare military orchid (Orchis militaris L.) at an established site in Suffolk, England

In Britain, where it reaches the north-westerly limit of its European distribution, Orchis militaris L. is extremely rare. Well-established and persistent populations of O. militaris are known to exist at only two sites. The largest extant population of O. militaris occurs in a disused chalk pit in Suffolk. A preliminary demographic analysis of this population, covering the period 1975 to 1991, along with estimates of key life stage transition probabilities are presented here. From 1975 to 1986 the number of separate identifiable plants in the population declined substantially. Until 1986 recruitment of rosettes was poor. The largest cohort of new plants, recorded in 1976, was 35. Approximately 48% of new individuals recruited between 1976 and 1985 failed to flower. Of those flowering, approximately 55% flowered during their first year above ground. Of the original population recorded in 1975, 67.8% flowered at least once during the study. The reproductive performance, i.e. the frequency of flowering and the period between episodes of flowering, varied considerably between individuals. Some plants flowered every year while others only flowered once during the study. Few plants remained below ground for more than one year, while several apparently persisted below ground for more than 6 successive years. Although the number of plants that can be identified as separate individuals has declined, the total number of rosettes in the population has, from 1986, increased dramatically. Because of the dense clumping of these recruits it is not possible to determine whether they are derived from seed or vegetative propagation. When post 1986 recruitment is combined with the number of plants that established before 1986 and survive, the apparent number of plants present at the site has more than doubled between 1975 and1991.     

Natural selection on gall size: variable contributions of individual host plants to population-wide patterns

Studies that provide estimates of the form and magnitude of selection on herbivore traits at the level of individual plants in natural populations represent a vital step in understanding the interaction of selection and gene flow among host-affiliated insect populations when individual plants equate to differing selective regimes. We analyzed phenotypic selection on the trait gall size for a host-specific gall former at both the individual host plant and population level (across host plants) in each of two years. Linear and nonlinear selection and the fitness function relating gall size to the probability of survivorship in the absence of natural enemies were estimated for each level and year. Selection imposed by the host plant was observed in 19 of the 22 subpopulations monitored. At the population level, linear and nonlinear selection were evident each year. However, population-level estimates masked the significant heterogeneity in the form and direction of selection evident among plants each year. Heterogeneity among gall-former subpopulations is emphasized by our findings that selection varied from directional to stabilizing among plants and the majority of selection gradients estimated for individual plants did not fall within the 95% CIs of the population-level estimates.     

Small‐scale resource tracking in a population of a long‐lived insect

How plant‐feeding insects distribute themselves and utilize their host plant resources is still poorly understood. Several processes may be involved, and their relative roles may vary with the spatial scale considered. Herein, we investigate small‐scale patterns, namely how population density of a gall midge is affected by individual growth form, phenology, and microsite characteristics of its herb host. The long‐lived plant individuals vary much with regard to number of shoots, flower abundance, and flowering phenology. This variation is connected to site characteristics, primarily the degree of sun exposure. The monophagous insect galls the flowers of the host plant – an easily defined food resource. It is a poor disperser, but very long‐lived; diapausing larvae can stay in the soil for many years. Galls were censused on individual plants during 5 years; from a peak to a low in gall population density. Only a very small fraction of the flowers produced (<0.5%) were galled even in the peak year. Nevertheless, most plant individuals had galls at least 1 year. In a stepwise multiple regression, plant size (number of shoots) was found to be the most important predictor of gall density (galls/flower). However, gall density decreased more than one order of magnitude over the plant size range observed. There was also a weak effect of plant phenology. Early flowering plants had lower gall densities than those starting later. Sun exposure had no direct effect on gall density, but a path analysis revealed indirect effects via the timing of flowering. Gall population change was highly synchronous in different parts of the study area with no significant decrease in synchrony with distance.     

Variation in the genetic structure and reproductive biology of holm oak populations

Five polymorphic enzymatic loci were studied in 30 populations of Quercus ilex distributed from Crete to North Africa. A high within-population genetic diversity was found for each locus studied. Most of the alleles were common to all the populations, indicating that the holm oak corresponds to a single genetic entity. Observation of breeding system characteristics showed that the holm oak is monoecious, wind-pollinated, and likely to possess genetic autoincompatibility. Furthermore, studies of flowering phenology in a single population showed that the period of flowering varied notably among trees. An average of 29% of the trees showed inter-annual variation in flowering time. The male, female or vegetative investment also varied widely among trees from one year to another. This results in a diversification of pollen source received by a given tree each year, and also from one year to another.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

Tree size and its relationship with flowering phenology and reproductive output in Wild Nutmeg trees

Reproductive strategies, sexual selection, and their relationship with the phenotype of individuals are topics widely studied in animals, but this information is less abundant for plants. Variability in flowering phenology among individuals has direct impact on their fitness, but how reproductive phenology is affected by the size of the individuals needs further study. We quantified the flowering intensity, length, and reproductive synchronization of two sympatric dioecious Wild Nutmeg tree species (Virola, Myristicaceae) in the Brazilian Atlantic forest, and analyzed its relationships with tree size. Two distinct strategies in flowering timing and intensity were found between species (annual versus biennial flowering), and among individuals in the annual flowering species (extended versus peak flowering). Only for the annual flowering species the reproductive output is related to tree size and large trees present proportionally higher flower coverage, and lower synchronization than smaller ones. Flowering is massive and highly synchronized in the biennial species. Sex ratios are not different from 1:1 in the two species, and in the two segregated reproductive subgroups in the biennial flowering species. The biennial flowering at individual level is a novelty among reproductive patterns in plants, separating the population in two reproductive subgroups. A proportional increase in the reproductive output with size exists only for the annual flowering species. A biennial flowering can allow resource storage favouring massive flowering for all the individuals diluting their relationship with size.     

The life history of Ranunculus glacialis, an arctic-alpine perennial herb, in Finnish Lapland

Dynamics and performance of established individuals of Ranunculus glacialis at the time of peak flowering were monitored in a mountain area in Finnish Lapland for five years (1982–1986). The plants reproduced only sexually, and their spatial distribution in 40 1-m² quadrats was clumped. During the study period the mean density rose from 8.5 to 10.3 individuals m⁻². The mean annual growth rate and birth rate of the population was 4% and 8.5%, respectively. The mean annual death rate of established individuals was 3.7%. The estimated mean life expectancy of an established individual was 27 yr. Death rate per quadrat was positively related to R. glacialis density, but birth rate per quadrat was negatively related to the percentage cover of stones per quadrat. High R. glacialis density per quadrat had negative effects, but high nitrogen availability had positive effects on the number of leaves per individual. An important factor reducing flowering was small rodent herbivory. In the same genets, the correlation between the number of leaves per individual in year t and the number of flowers per individual in year t+4 was significant. The below-ground biomass of R. glacialis comprised about 70% of the total biomass, and there was a close correlation between the above- and the below-ground biomass of an individual. The individuals flowered abundantly each year and produced a great number of small-sized achenes.     

Inter-populational variation,but no-annual variation within populations,in terms of reproductive size and genetic structure in a monocarpic perennial herb,Cardiocrinum cordatum var. glehnii

Cardiocrinum cordatum var. glehnii (Liliaceae) is a monocarpic perennial herb living in temperate broad-leaved deciduous forests. In the present study we examined the sizes (basal diameter of the stem and flower numbers) of flowering individuals and genetic diversity using microsatellite loci of 23 populations of C. cordatum var. glehnii in Hokkaido, Japan, over 2 years (2009 and 2010). As a result, we found both the basal stem diameter and the number of flowers varied widely among the populations. However, although the sizes of flowering individuals differed among the populations, these were very stable in each population and in each year. In addition, for genetic diversity, the same trends (i.e. wide variation among the populations but non-annual variation) were detected.     

Effects of local density on insect visitation and fertilization success in the narrow-endemic Centaurea corymbosa (Asteraceae)

Plant population size and density can influence the interactions between plants and pollinators, and affect plant reproductive success. We investigated the effects of local conspecific density on insect visitation and fertilization success in the rare, cliff-dwelling, self-incompatible Centaurea corymbosa , in which fecundity plays a key role in variation in population growth rate among years and among the six extant populations. Plant size, capitulum size, the abundance of co-flowering species, and the weather conditions were added as covariates in the analyses. Over three populations and two years (1995 and 2002), fertilization rate was positively related to the number of flowering conspecifics within 10 m. Fertilization rate varied among populations, but this variation over all six populations in 2002 could not be attributed to differences in population size. Observations in one population in 2003 showed that there was no difference in insect visitation per capitulum between plants in sparse vs dense patches whilst plants from sparse patches suffered reduced fertilization rate. Visitation and fertilization rates were not affected by plant size and the abundance of co-flowering species, but weather conditions at the time of flowering had a strong effect on both variables. Capitulum size had a positive effect on visitation rate, but an effect on fertilization rate only in 1995 and 2002 and not in 2003. Our results suggest that pollen limitation affects fertilization rate in C. corymbosa due to limited compatible mate availability rather than pollinator limitation. They agree with previous genetic results derived from paternity analysis. Whether or not the benefits of local spatial agregation to reproductive success result in increased individual fitness will depend on the relative reduction in survival of vegetative stages due to intra-specific competition.