扎龙湿地芦苇分株生态可塑性及其对土壤因子的响应

扎龙湿地的芦苇既可形成大面积的单优群落,也可形成不同群落斑块。采用大样本抽样调查与统计分析方法,对湿地内水生生境、湿生生境、旱生生境和盐碱生境芦苇种群分株高度和生物量进行比较。结果表明,6—10月份,4个生境芦苇种群分株高度及生物量均以水生生境最高,盐碱生境最低,水生生境株高为盐碱生境的1.5—2.3倍,分株生物量为2.0—5.1倍,生境间的差异性以及差异序位均相对稳定。4个生境株高生境间变异系数(19.45%—31.56%)均高于生境内变异系数(8.07%—17.61%),分株高度在生境间的可塑性更大;分株生物量中水生生境、湿生生境和盐碱生境3个生境间的变异系数(33.43%—55.61%)均低于生境内变异系数(44.85%—79.82%),分株生物量在生境内的可塑性更大。不同生境条件下芦苇种群分株,在生长和生产上均存在较大的生态可塑性,表现出明显的环境效应,其中土壤含水量是该地区芦苇分株生态可塑性变异的主要驱动因子(R0.80),为正向驱动。     

扎龙湿地不同生境芦苇分株的形态特征及生物量分配的异速关系

不同环境条件下的植物个体可以表现出形态特征的分异和物质分配的权衡与调整。采用大样本抽样调查与统计分析方法,比较研究扎龙湿地不同生境芦苇(Phragmites Australis)生殖株和营养株的形态特征以及生物量分配的异速关系。结果表明：在9月末,盐碱生境、旱生生境、湿生生境和水生生境芦苇分株的生长表现出较大的生态可塑性,株高和株重均以盐碱生境最小,水生生境最大,最大值与最小值的比值分别为1.3—3.3和1.8—5.1,分株生长在种群间的变异度高于种群内,与营养株相比,生殖株的变异度较低;分株的支持分配与生产分配的比值为1.8—4.2,生产分配以盐碱生境最高,以水生生境最低,而支持分配和生殖分配表现与生产分配相反的序位;生殖株的花序长和花序重与株高间呈直线函数形式增长,株高和株重低于种群平均值的20%和35%的分株不进行有性生殖;叶重、叶鞘和茎重以及分株重与株高间呈幂函数形式的异速生长关系。植物通过改变个体的形态特征以及调整构件间生物量分配适应不同环境,而受遗传因素控制的构件间生长关系却相对稳定。     

Production of male flowers does not decrease with plant size in insect‐pollinated Sagittaria trifolia,contrary to predictions of size‐dependent sex allocation

Abstract It has been proposed that relative allocation to female function increases with plant size in animal‐pollinated species. Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size‐dependent sex allocation (SDS), throwing doubt on the generalization of SDS. Plant size, phenotypic gender, and flower production were measured in experimental populations of an aquatic, insect‐pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities. The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors. In the high‐density population, 48% of ramets were male without female flowers, but in the low‐density population all ramets were monoecious. We observed allometric growth in reproductive allocation with ramet size, as evident in biomass of reproductive structures and number of flowers. However, within both populations female and male flower production were isometric with ramet size, in contrast to an allometric growth in femaleness as predicted by SDS. Phenotypic gender was not related to ramet size in either population. The results indicated that large plants may increase both female and male function even in animal‐pollinated plants, pointing towards further studies to test the hypothesis of size‐dependent sex allocation using different allocation currencies.     

扎龙湿地保护区异质生境芦苇种群分株构件的数量特征

采用单位面积取样,计数和测量的调查与统计方法,对扎龙湿地保护区4个生境单优群落芦苇种群分株构件数量特征进行比较分析。结果表明,4个生境芦苇种群从5月10日左右返青后进入营养生长期,分株高度和分株密度均持续增加到生殖生长初期的8月份,其中6-8月份差异均达到显著水平(P0.05),8-10月份差异均未达到显著水平(P0.05),芦苇进入生殖生长期后,分株便停止高度生长,地下芽的输出也不再形成分株补充现实种群;分株生物量和种群生物量均持续增加到生殖生长旺盛期的9月份,至休眠期的10月份均有所降低,各月份间的差异均达到显著水平,芦苇种群在生长季末期,具有将生产的物质分配转移到地下储藏与营养繁殖器官的形成与生长上的特性。芦苇种群分株数量特征与返青后实际生长时间之间均较好地符合对数函数关系,R~2在0.818-0.994之间,拟合方程均达到了P0.05的显著水平。4个生境芦苇种群分株密度以湿生生境最大,依次为水生生境、旱生生境,盐碱生境最小,分株高度、分株生物量和种群生物量均以水生生境最大,依次为湿生生境、旱生生境,盐碱生境最小。因此,4个生境芦苇种群分株构件数量特征均表现出基本一致的生育期节律性,同时,芦苇种群的个体生长和种群动态存在明显的环境效应,土壤含水量、pH是影响该地区芦苇分株数量特征的主要因子。     

Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation

It has been proposed that relative allocation to female function increases with plant size in animalpollinated species.Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size-dependent sex allocation (SDS),throwing doubt on the generalization of SDS.Plant size,phenotypic gender,and flower production were measured in experimental populations of an aquatic,insect-pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities.The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors.In the high-density population,48％ of ramets were male without female flowers,but in the low-density population all ramets were monoecious.We observed allometric growth in reproductive allocation with ramet size,as evident in biomass of reproductive structures and number of flowers.However,within both populations female and male flower production were isometric with ramet size,in contrast to an allometric growth in femaleness as predicted by SDS.Phenotypic gender was not related to ramet size in either population.The results indicated that large plants may increase both female and male function even in animal-pollinated plants,pointing towards further studies to test the hypothesis of size-dependent sex allocation using different allocation currencies.     

Life-history monographs of Japanese plants 14: Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae)

The life-history characteristics of Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae) are described. The species is an herbaceous perennial that favors open but relatively moist habitats. It is distributed from Kyushu to Hokkaido in Japan but also occurs in Korea, Mongolia, China and the Russian Far East. In southwestern Japan, shoots start sprouting from mid- to late April, reaching approximately 30–70 cm in height, with 5–10 compound leaves. Ramets simultaneously produce one or more stoloniferous rhizomes that elongate until new ramets are formed at the tips. Cardamine leucantha has a pseudo-annual life cycle, in which mother ramets wither at the end of each season and only daughter ramets appear aboveground in the next year. As a result, ramet positions change annually. In a study population, the number of flowers averaged 23.9 ± 21.0 per ramet and fruit set was 44.2 ± 24.8% (10.4 ± 10.1 fruits per ramet). Ramets produced 3.8 ± 2.3 rhizomes that were 22.0 ± 15.6 cm long. The species sometimes forms large populations. A single genet develops into a group of disconnected ramets spreading via clonal growth. Reproductive characteristics (e.g., fruit set and numbers of flowers and rhizomes) vary among populations, resulting in interpopulation differences in genet structure. Because the reference genome became available recently, established molecular tools will be applied effectively for the investigations of C. leucantha as a model clonal plant.     

Factors controlling the population dynamics of the clonal helophyte Ranunculus lingua

Abstract. Two marginal and two central populations of the pseudo-annual aquatic plant Ranunculus lingua were studied over four years. The main purpose was to quantify potentially influential abiotic and biotic factors and to derive predictions about life-history differences between the populations. Variation in abundance and height of R. lingua ramets at different depths were related to water-level fluctuations, to abundance of other helophyes (emergent macrophytes), and to the occurrence of invertebrate grazing and fungal pathogens. Clear differences between marginal and central populations were shown in the depth distribution of ramet numbers and ramet heights, as well as in the dynamic patterns, where marginal populations had a higher flux of ramets. These patterns and regression analyses indicated that abiotic factors have a greater influence in marginal populations, whereas biotic factors are more important in central populations. It is suggested that marginal habitats for R. lingua would favour life-histories with a high reproductive capacity, whereas a large size of ramet would be the most important life-history feature in central habitats. This was supported by the fact that ramets in marginal populations, in spite of their smaller size, produced higher number of rhizomes than ramets in central populations. Variation in regional abundance was finally related to differences in demographic processes and dispersal potential between the populations.     

松嫩平原不同生境芦苇种群分株的生物量分配与生长分析

在松嫩平原8月中旬,4个旱地生境芦苇种群分株的生长和生产力以季节性临时积水的低洼地为最高,以碱斑地为最低,两个生境之间分株高度相差2.8倍,分株生物量相差4.4倍,其它2个生境沙地和草甸混生 少也达到了显著或极显著水平,不同生境芦苇种群分株生物量分配的差异,蕴涵着重要的生长调节和物质分配策略。在分株株较小而又有充分生长空间的沙地生境和三斑生境,以及具有种间竞争的草句生境,分株将50％以上的物质分配给叶的建造上,以保证充分的物质生产;在环境条件相对较好的低洼地生境,分株将近40％的物质分配给茎的伸长生长,以提高个体对空间和光资源的竞争力。4个生境芦苇种群在分株增高与增重,以及茎、叶与分株的相对增重均具有相同的幂函数异速生长规律。     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

Evolution of an invasive phenotype: shift to belowground dominance and enhanced competitive ability in the introduced range

In response to novel selection pressures in an introduced range, non-native species may evolve more competitive phenotypes unique from those of their native range. We examined the existence of an invasive phenotype in the herbaceous perennial Artemisia vulgaris, a frequent invader of the Northeast and Mid-Atlantic US. Populations from both the native (European) and the introduced (North American) ranges were grown in intra-specific competition (same population), inter-specific competition with the native perennial herb Solidago canadensis, and alone in a common garden to quantify shifts in resource allocation and neighbor effects on performance and competitive ability. Without competition, introduced A. vulgaris populations were much shorter than native populations, but germinated earlier, produced more ramets, more belowground and total biomass, and maintained higher root-to-shoot ratios. Under inter- and intra-specific competitions, introduced A. vulgaris populations were shorter, but produced more ramets, belowground, and total biomass than native populations. S. canadensis belowground and total biomass were more highly suppressed by introduced than native A. vulgaris. Our data suggest that since the introduction to North America, A. vulgaris has evolved a more competitive invasive phenotype characterized by many short ramets with more extensive root/rhizome networks. This rapid evolutionary shift likely benefits A. vulgaris in its introduced range by allowing establishment and subsequent dominance in dense stands of existing vegetation.     

黄帚橐吾花序结构的资源配置与环境的关系

 为了系统地了解黄帚橐吾(Ligularia virgaurea) 自然种群的繁殖对策,在其繁殖分配研究的基础上,通过统计不同部位头状花序的生物量投入,进一步分析了存在于总状花序内资源分配上的结构效应及其对不同生境条件的反应。结果表明：1）总状花序、不同部位头状花序在资源投入上受其栖息生境各种生态因子的影响而在不同生境和平均种群密度间存在着差异,并表现出一定的趋势;2）可代表个体水平的总状花序大小、头状花序大小、头状花序数量和头状花序平均重量等特征,都与种群密度呈现出程度不同的负相关关系,而总状花序内不同部位的头状花序大小则与种群密度变化无关;3）总状花序内的资源分配存在着位置依赖性,这种显著差异表现为顶部头状花序比基部和中部头状花序有较大的资源分配;4）总状花序顶部、中部和基部头状花序的资源分配与总状花序大小表现出极显著或显著的负相关关系。     

Impact of soil water chemistry on the apparent sex ratio of the flowering ramets of the dioecious plant Myrica gale var. tomentosa

We determined whether the apparent (M/Fl) sex ratio (male ramets/flowering ramets) and apparent reproductive ramet ratio (Fl/Li ratio; flowering ramets/living ramets) in 15 Myrica gale var. tomentosa populations varied with dissolved total nitrogen, dissolved total phosphorus, potassium, magnesium, calcium, or pH in the soil water. Our aim was to define the environmental factors affecting the M/Fl sex ratio and Fl/Li ratio of the populations. We also examined the habitat conditions of these populations by analyzing soil water chemistry and water dynamics. In 2007, 3 of the 15 populations had no females. The remaining 12 had significantly male-biased (M/Fl sex ratio?=?0.59-0.97). Although we could not explain the absence of females by the current potassium levels alone, as potassium increased, so did the M/Fl sex ratio. As nitrogen increased and potassium decreased, Fl/Li ratio decreased. Our soil water chemistry analyses suggested that the potassium supply by soil surface erosion from flooding and the inflow of anthropogenic nitrogen were the important factors influencing the M/Fl sex ratio and Fl/Li ratio. Nitrogen management would be important in one of the endangered populations where inflow of nitrogen was the highest among 15 habitats.     

A predictive relationship between population and genetic sex ratios in clonal species

Sexual reproduction depends on mate availability that is reflected by local sex ratios. In species where both sexes can clonally expand, the population sex ratio describes the proportion of males, including clonally derived individuals (ramets) in addition to sexually produced individuals (genets). In contrast to population sex ratio that accounts for the overall abundance of the sexes, the genetic sex ratio reflects the relative abundance of genetically unique mates, which is critical in predicting effective population size but is difficult to estimate in the field. While an intuitive positive relationship between population (ramet) sex ratio and genetic (genet) sex ratio is expected, an explicit relationship is unknown. In this study, we determined a mathematical expression in the form of a hyperbola that encompasses a linear to a nonlinear positive relationship between ramet and genet sex ratios. As expected when both sexes clonally have equal number of ramets per genet both sex ratios are identical, and thus ramet sex ratio becomes a linear function of genet sex ratio. Conversely, if sex differences in ramet number occur, this mathematical relationship becomes nonlinear and a discrepancy between the sex ratios amplifies from extreme sex ratios values towards intermediate values. We evaluated our predictions with empirical data that simultaneously quantified ramet and genet sex ratios in populations of several species. We found that the data support the predicted positive nonlinear relationship, indicating sex differences in ramet number across populations. However, some data may also fit the null model, which suggests that sex differences in ramet number were not extensive, or the number of populations was too small to capture the curvature of the nonlinear relationship. Data with lack of fit suggest the presence of factors capable of weakening the positive relationship between the sex ratios. Advantages of this model include predicting genet sex ratio using population sex ratios given known sex differences in ramet number, and detecting sex differences in ramet number among populations.     

EFFECTS OF FIRE ON CLONAL GROWTH AND DYNAMICS OF PITYOPSIS GRAMINIFOLIA (ASTERACEAE)

The rhizomatous perennial Pityopsis graminifolia was studied in a Florida sandhill community in an annually burned site, a periodically burned site, and a site that has been protected from fire since 1965. These different fire regimes significantly affected the demography and life histories of both plants and plant parts in this clonal species. Fires resulted in reductions in ramet biomass and height, and an increase in the (root + rhizome)/shoot biomass ratio. Burning also decreased the total number of flower heads and new rhizomes produced per ramet. However, the survivorship of initiated rhizomes was greater in burned sites and resulted in a larger number of established daughter ramets per clone. As a result, in burned sites there was a shift in clone structure toward larger numbers of smaller ramets, but there were no significant reductions in seed or rhizome production on a per genet basis. The results showed that the responses to fire in P. graminifolia are different when measured at the genet vs. ramet level and that the effects of fire on clones can be explained by demographic responses of plant parts. Population regeneration in the study sites was dependent on successful clonal ramet production because no seedling recruitment was observed. This suggests that disturbances other than fire are important for new genet recruitment in these clonal populations.     

Ramet demography and ecological attributes of the perennial river corridor plant Cnidium dubium (Schkuhr) Thell. (Apiaceae)

Cnidium dubium is a highly endangered, perennial river corridor plant in Central Europe. Here we summarise a 4-year field study of its ramet demography, growing in two different flood meadows in the Lower Havel River Valley in northeastern Germany, emphasising the influence of site-specific abiotic factors, mowing and the summer flood of 2002.We recorded 1658 ramets at densities between 2 and 180 ramets m⁻², which varied over area and time, indicating a high small-scale turnover of this species within grasslands. The half-life of the different ramet cohorts was estimated to range from 0.1 to 1.3 years, but single ramets of more than 4 years old were also found. Ramets are monocarpic: they propagate only once in their life. The highest probability of a vegetative ramet becoming a flowering stalk was 11.6%. Ramet density, cumulative number of newly produced ramets per year, leaf length, leaf age, reproductive growth and recovery from the summer flood of 2002 varied, depending on site. C. dubium had a higher performance at sites in intermediate zones of the overall flooding gradient compared with higher zones. Flooding stress is the controlling factor for the lower limit of its performance, while there was no evidence for an upper limit controlled by summer drought. Mowing slightly increased the cumulative number of leaves per ramet, but it decreased the size of new leaves and also the summer above-ground biomass per ramet. As a consequence, the most marked effect was that fewer ramets entered the generative life stage, but the plants were able to keep their overall reproductive performance on a level independent of mowing. Knowledge of basic growth patterns of C. dubium in natural populations could help to explain the species rareness, and may also suggest appropriate strategies to manage its survival.     

Distylic Hottonia palustris shows high reproductive success in small populations despite low genetic variability

Hottonia palustris L. (Primulaceae) is characterized by a heteromorphic incompatibility system. The strategy of distylic ramets is believed to promote outcrossing, to maintain overall genetic diversity and to prevent inbreeding depression. In spite of this distyly, an extremely low amount of allozymic diversity was observed in 545 individual ramets from 14 populations in Flanders (Belgium). A possible explanation for such low genetic variation is discussed in relation to the vegetative propagating abilities and the ecological niche width of the species. In contrast to the uniformity in allozymes as well as to the feature of single morph populations, there was a high variability in reproductive success between populations such as the number of seeds per ramet (425–2633), the number of flowers per ramet (9–36) and the mean weight of seeds (0.03–0.17 mg). Small populations and even those consisting of only one style morph may show a high reproductive success. As a whole, H. palustris showed a negative relationship of reproductive success with the surface area of its populations.     

Differential costs of sexual and vegetative reproduction in wild strawberry populations

Summary The CO₂ costs of producing sexual and vegetative reproductive propagules were calculated for two species of wild strawberry, Fragaria virginiana and F. vesca. Five populations on sites representing a gradient of successional regrowth near Ithaca, New York, USA, were studied for two or three years each. Field studies of phenology, biomass, demography, and environment and laboratory studies of CO₂ exchange were integrated using a computerbased model of CO₂ dynamics to estimate costs of propagules.The percentage of plants flowering and the number of flower buds produced were highest in an open, recently disturbed habitat and lowest in a forest habitat. The openhabitat plants had the greatest success in converting flower buds into ripe fruits and also produced the highest numbers of runners and runner plantlets. On the basis of total investments in structure and respiration minus any photosynthetic gain of all reproductive structures, the cost per seed was lowest in the most open habitats and highest and increasingly variable in the more closed habitats. The cost of plantlets also was lowest in the most open habitat. The differences among habitats in cost of plantlets alive after one or two growing seasons increased due to differential survivorship of plantlets, with the open habitat continuing to have the lowest cost per plantlet. Theoretical treatments of life history characteristics such as reproductive effort should recognize that costs of equivalent type and size of propagule may vary among environments.     

Do clonal growth form and habitat origin affect resource-induced plasticity in Tibetan alpine herbs?

To investigate how growth form and habitat origin affect phenotypic plasticity to resource supply in the Tibetan alpine herbs, the phalanx-type species Stipa capillacea and the guerilla-type species Carex montis-everestii were sampled from two different habitats (alpine steppe and alpine scrubland) and grown under three levels of light intensity and two levels of nutrient supply. Interspecific differences in light-induced plasticity were detected only in number of ramets, specific leaf area and leaf sheath length. Plasticity in plant biomass, number of ramets and rhizome length in response to light intensity differed between the two habitats. Stipa plants were more plastic than Carex plants in number of ramets and specific leaf area in response to light intensity. Carex plants from the alpine scrubland expressed greater light-induced plasticity in plant biomass and number of ramets than those from the alpine steppe, and Stipa plants showed less interhabitat differences in plasticity, which may be closely related to their contrasting growth forms. Clonal growth form and habitat origin affected nutrient-induced plasticity in none of the measured traits. It may be the guerilla growth form that makes Carex plants more efficiently adapted to highly heterogeneous light conditions in scrubland, and less habitat-dependent plasticity contributes to success of the phalanx-type Stipa plants in alpine habitats. The results are discussed in the context of foraging for heterogeneously distributed essential resources and adaptation to habitat origin.     

Growth behaviour of the invasive species <Emphasis Type="Italic">Ipomoea carnea</Emphasis> in the Nile Delta,Egypt

Ipomoea carnea Jacq., a native shrub of South America, grows in dense populations along river beds, river banks, canals and other waterlogged (wetland) areas. It has become naturalised along canals, drains, road sides and field edges in the Nile Delta, Egypt. The rapid growth rate, spread and adaptability from xeric to aquatic habitats indicates that this plant may potentially become a disastrous invasive species in Egyptian water bodies. Fifty established stands covering seven habitats (railway sides, waste lands, road sides, drain and canal banks, road dividers and field edges) in the Nile Delta were used in this experiment. In each stand, 20 ramets were permanently marked to estimate the monthly variations in such growth variables as phenology, height and diameter of the ramet canopy, number of flowers, leaves and fruits. The results showed a significant variation in growth variables relative to habitat types. Generally, I. carnea populations along the railway sides and field edges had the lowest values for most variables; while those of the road dividers, road sides and canal banks produced the highest values. The growth of I. carnea follows a seasonal pattern where the highest values for most growth variables were obtained during September and October and the lowest were recorded during June and July. The greatest flower and fruit production was observed between September and December. Similarly, it was shown that the flowering time varied in relation to habitat types; it began earlier on road sides and canal banks, but later on railway sides, road dividers and waste lands. Leaf areas of I. carnea populations in the wet habitats (e.g. drain and canal banks) were higher than those in other habitats. These results are discussed in relation to other related studies.     

Influence of local floral density and sex ratio on pollen receipt and seed output: empirical and experimental results in dichogamous Alstroemeria aurea (Alstroemeriaceae)

Local density and sexual composition are two aspects of floral neighborhoods thought to influence pollination and seed output of recipient plants. I characterized the floral neighborhood of 436 flowering ramets of Alstroemeria aurea, a southern Andean perennial, distributed among three sites. On each ramet, I measured total pollen receipt and seed output. The long-lived, bumblebee-pollinated flowers of A. aurea are synchronously protandrous with a given ramet being either all male or all female and thus incapable of self or geitonogamous pollination at the ramet level. Even though each ramet changes sex over time, A. aurea forms floral neighborhoods that remain stable with respect to density and sex ratio during the span of a focal ramet female phase. Contrary to expectation, under field conditions neither local density nor sexual identity explained significant amounts of variation in pollen receipt. Density of neighboring flowering ramets marginally affected pollen receipt in two of the three populations but in opposite directions. Despite the absence of strong effects of neighborhood sexual composition on pollen receipt, the sexual identity of neighbors affected seed output which suggests effects on the quality of pollination due to changes in patterns of pollen flow. I also compared pollen loads on the stigmas of artificially isolated ramets (6 m) with those on experimental focal ramets surrounded by six close neighbors (20 cm) that were either all male or all female. Here, pollen receipt by focal ramets in all-male neighborhoods was 1.3 times greater than in isolated ramets, and 3.8 times greater than in ramets in all-female neighborhoods. In these artificial neighborhoods, stigmatic pollen deposition increased significantly over time. In nature, rates of bumblebee visits were higher in female-biased (early-flowering) than in male-biased (late-flowering) co-occurring floral patches. Thus, spatio-temporal shifts in visitation frequencies associated with the sexual composition of floral neighborhoods might compensate for spatial variability in pollen availability within populations and explain the discrepancies between empirical and experimental results. Received: 11 June 1996 / Accepted: 24 March 1997