Dissecting components of flowering pattern: size effects on female fitness

Flowering synchrony is essential for plant reproductive success, especially in the case of small‐sized populations of self‐incompatible species. Closely related to synchrony, flowering intensity influences pollinator attraction and pollinator movements. Thus, a high flowering intensity may increase pollinator attraction but, at the same time, may also increase the probability of geitonogamous pollinations. Depending on the mating system, the female fitness of plants in small populations may be affected by both the positive effects of higher flowering synchrony and pollinator attraction and the negative effects of geitonogamous pollinations induced by a high flowering intensity. It was hypothesized that different‐sized plants in a population would show contrasting flowering patterns, resulting in differences in pollinator behaviour. These influences could result in differences in mating and female reproductive success. This hypothesis was tested by studying the flowering pattern of Erodium paularense (Geraniaceae), a rare and endangered endemic of central Spain. The temporal distribution of flower production was explored throughout the reproductive season, and the probability of xenogamy and geitonogamy and their relationship to plant size and fitness components were calculated. The analysis of this partially self‐compatible species showed diverse flowering patterns related to different plant sizes. Small plants produced a larger number of seeds per fruit in spite of having lower values of flowering synchrony. By contrast, large plants produced a larger number of seeds from geitonogamous pollinations. The effect of different flower displays and outcrossing rates on seed set varied throughout the season in the different groups. Our findings highlight the relevance of individual plant size‐dependent phenology on female reproductive success and, in particular, on the relationship between flowering synchrony and fitness. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156 , 227–236.     

Consequences of variation in flowering phenology for seed head herbivory and reproductive success in Erigeron glaucus (Compositae)

Summary We examined the relationship between flowering phenology, reproductive success (seed production only), and seed head herbivory for 20 similarly sized clones of Erigeron glaucus growing at Bodega Bay Reserve, northern California, USA. Although clones tended to reach peak flowering on the same date, they differed in the proportion of their total flowers produced around that date (flowering synchrony). Clones also differed in the number and density of flower heads presented at any one time to pollinators and herbivores (floral display). Both of these characteristics had consequences for herbivory and plant reproductive success. The proportion of flower heads damaged by insect herbivores was greater for clones that concentrated flowering activity during the main flowering period for the population as a whole (high synchrony) compared to clones that spread flowering out temporally. The primary reason for this result was that clones with low flowering synchrony produced a significant proportion of their flower heads during the fall and therefore, escaped attack by the tephritid fly, Tephritis ovatipennis. Clones with intermediate synchrony had lower seed success (total number of viable seeds produced over the year) than clones with either low or high synchrony. The proportion of flower heads damaged by insect herbivores and number of tephritid flies reared from flower heads were both negatively correlated to floral display while seed head mass and germination rates were positively related to display. Thus, clones which produced dense floral displays were favored both in terms of reduced herbivory and increased successful seed production.     

Effects of plant size and weather on the flowering phenology of the organ pipe cactus (Stenocereus thurberi)

Background and Aims

Flowering phenology is a critical life-history trait that influences reproductive success. It has been shown that genetic, climatic and other factors such as plant size affect the timing of flowering and its duration. The spatial and temporal variation in the reproductive phenology of the columnar cactus Stenocereus thurberi and its association with plant size and environmental cues was studied.

Methods

Flowering was monitored during 3 years in three populations of S. thurberi along a latitudinal gradient. Plant size was related to phenological parameters. The actual and past weather were used for each site and year to investigate the environmental correlates of flowering.

Key Results

There was significant variation in the timing of flowering within and among populations. Flowering lasted 4 months in the southern population and only 2 months in the northern population. A single flowering peak was evident in each population, but ocurred at different times. Large plants produced more flowers, and bloomed earlier and for a longer period than small plants. Population synchrony increased as the mean duration of flowering per individual decreased. The onset of flowering is primarily related to the variance in winter minimum temperatures and the duration to the autumn–winter mean maximum temperature, whereas spring mean maximum temperature is best correlated with synchrony.

Conclusions

Plant size affects individual plant fecundity as well as flowering time. Thus the population structure strongly affects flowering phenology. Indications of clinal variation in the timing of flowering and reproductive effort suggest selection pressures related to the arrival of migrating pollinators, climate and resource economy in a desert environment. These pressures are likely to be relaxed in populations where individual plants can attain large sizes.Key words: Flowering phenology, optimal timing, plant size, Sonoran Desert, Stenocereus thurberi, temperature     

准噶尔无叶豆的开花物候与生殖特征

 为了研究沙漠稀有植物准噶尔无叶豆(Eremosparton songoricum)的开花物候特征及其对生殖成功的影响, 2005和2006年连续两年对其自然种群的开花物候和开花过程中的花部表型变化进行了观察, 并运用相对开花振幅、开花强度和开花同步性等开花物候指数研究了开花物候特征。结果表明: 准噶尔无叶豆在5月下旬至6月中下旬开花, 其种群、个体、花序和单花的花期分别历时26~29 d、8~10 d、5~7 d和2~3 d。单花开花进程依其形态和散粉特征可分为散粉前期、散粉初期、散粉盛期和凋谢期4个时期。其个体水平的开花物候进程(开花振幅曲线)呈渐进式单峰曲线, 具有很高的开花同步指数, 表现出一种集中开花的模式。开花物候指数与座果数之间的相关分析结果表明, 始花日期与花期持续时间存在负相关关系, 而与开花数和座果数存在正相关关系; 花期持续时间与开花数和座果数存在显著正相关关系。准噶尔无叶豆个体开花物候在很大程度上是由其遗传因子决定的, 而开花物候在年度间的变异, 可能是由于荒漠气候的差异(主要是水分和温度的差异)所引起的。作为沙漠窄域分布特有种, 准噶尔无叶豆在环境和人为干扰的双重选择压力下, 为了吸引更多的传粉者访问而达到生殖成功, 形成了大量集中开花的模式。     

Effects of population size on reproductive success of the endangered and endemic species Primula merrilliana

The reproductive ability related to the population size of the endangered and endemic species Primula merrilliana Schltr.was investigated.In 26 populations observed,only four contain more than 500 flowering individuals,whereas most of them(53.8%) consist of less than 100 flowering individuals.Though the ratio of pin and thrum plants keeps its balance at 1:1 for all populations,the frequency of pin and thrum flowers was significantly biased in most small populations.Population size strongly affected reproductive success; plants in small populations produced significantly fewer flowers and fruits per plant and fewer seeds per fruit,and therefore fewer seeds per plant.The floral density was another important factor that influenced the reproductive success of P.merrilliana,because four main reproductive success parameters (fruits per plant,seeds per fruit,seeds per plant,and the proportion of flowers setting fruit) were all positively correlated with floral density.The size of plants and the number of leaves per plant (measure of habitat quality) were not influenced by the variation of population size,suggesting that the reduced fecundity in small populations may not be a consequence of lower habitat quality,inbreeding depression and pollen limitation as a result of less attractiveness in small populations are therefore likely explanations for these patterns.     

荒漠孑遗植物裸果木的开花物候特征

2014年和2015年连续2年对哈密和乌恰裸果木(Gymnocarpos przewalskii)自然种群的开花物候特征进行野外定点观测,并运用相对开花振幅、开花强度和开花同步性等参数对其开花物候特征进行分析。结果表明,2个种群每年均开1次花,4月下旬到5月上旬进入始花期,开花进程基本相似,均呈单峰曲线,具有较高的开花同步性,呈现"集中大量"开花模式;在个体水平上,始花期、终花期在种群和年份间略有差异;单花持续时间在年际间是稳定的,但在种群间有所不同,即乌恰种群的单花持续时间略长于哈密种群;不同裸果木种群的开花数与结籽数间均呈极显著正相关,开花同步性指数与相对开花强度间呈显著负相关;乌恰种群的始花期与开花数间呈负相关。裸果木个体开花振幅、开花同步指数及开花模式在种群和年份间的相似性很大程度上是由其在长期的地理变迁中形成的遗传特性决定的,而始花期和开花持续时间等开花物候特征的差异主要是由环境和气象因子(海拔、气温、降水及光照)差异造成。研究结果对于探讨该属植物的繁殖生物学特性及其保护具有重要意义。     

VARIATION IN INDIVIDUAL FLOWERING TIME AND REPRODUCTIVE SUCCESS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Field studies on two populations of Agalinis strictifolia were conducted over a 3-year period to investigate the relationship between flowering time of individuals and plant size, flowering duration, flower and fruit production, fruit predation, and growth rate. Seasonal patterns of pollinator visitation were compared with those of individual flowering time, flower density, percent fruit production, and mean seeds/fruit. In general, early and middle flowering individuals (as determined by either first flowering date or peak flowering) were larger, flowered longer, and produced more flowers and fruits than late flowering individuals. Early and middle flowering individuals (based on first flowering date) also grew faster than late flowering individuals. Although early and middle flowering individuals produced more fruits, fruit predators did not damage a disproportionate number of fruits compared to late flowering individuals. Patterns of bee visitation showed no association with seasonal patterns of flower density, percent fruit production, mean seed/fruit, or individual flowering time. In populations of A. strictifolia, it would seem that biotic or environmental determinants of growth rate (hence size and reproductive success) may be more important in generating variation in individual flowering time than patterns of pollinator visitation or fruit predation.     

濒危植物长柄双花木开花物候与生殖特性

研究了长柄双花木开花过程中花部表型的变化。连续4a对其野生种群、1a对人工种群的开花物候进行了观察,并运用相对开花强度和同步性等开花物候指数分析了开花物候对其生殖的影响。结果如下：长柄双花木开花时间为9月上、中旬至11月中、下旬;单花花期一般为6～7d,单花依其形态和散粉特征可以分为4个时期：散粉前期、散粉初期、散粉盛期和凋谢期。个体开花持续时间49～55d,种群花期历时63～71d。种群内不同年度间开花物候指数没有显著差异,而种群间则存在显著差异,野生种群开花进程为渐进式单峰曲线。人工种群则为“钟”形曲线,二者均属于“集中开花模式”。长柄双花木具有2个相对开花强度的分异趋势,这种分异趋势具有进化意义。开花物候指数与生殖间的相关分析表明,始花时间与开花数量、座果率及花期长度之间均具显著负相关关系,而开花数量与花期长度之间则呈显著正相关,但均为线性相关。长柄双花木开花物候在种群间的差异和种群内年度间的相似性说明,其开花时间可能是由与其相关的复杂的微生境特征和(或)由其遗传因子决定的,同时也反映了种群间的遗传分异和种群内个体间的遗传一致性。作为一种濒危物种,长柄双花木在这种环境的选择压力之下,形成了“大量、集中开放的花”的开花模式,吸引到更多的传粉者的访问,从而达到生殖成功。     

Relationships between flowering phenology,plant size and reproductive success in shape Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.     

濒危植物长柄双花木自然种群生殖构件的时空动态

根据野外观测数据,研究了濒危植物长柄双花木(Disanthus cercidifolius var.longipes)在不同生境条件下自然种群开花物候变异以及生殖构件的时空动态.结果表明,不同群落类型中长柄双花木开花物候存在一定差异,在纯林中个体的平均花期最长,达75d,而个体间开花同步性最低,仅为0.717;竹林中的个体花期最短,只有47d,而同步指数最高,达0.968.生殖枝数量及花序数均随个体年龄增长而增加,在年龄为30～35年时达最高,此后又呈下降趋势;在海拔为810m左右处,个体生殖枝数及着生的花序数均达最大值,且冠层间差异明显.纯林中个体着生的生殖枝数及每生殖枝上着生的花序数均最高,分别为411.39和7.857;竹林中个体每生殖枝着生的花序数显著低于其它群落.年龄小的个体,各时期生殖构件败育率均高于年龄大的个体;另外,生殖构件的败育与海拔高度间无相关性,而与构件发育时间、分布的树冠层次以及所处的群落类型相关.影响生殖构件败育的主要因子是光照条件.     

Plant size,flowering synchrony and edge effects: What,how and where they affect the reproductive success of a Neotropical tree species

Plant reproductive success is supposedly influenced by phenology and individual size, which may be modified under edge effects. We tested if reproductive success, estimated by fruit set, in Senefeldera verticillata (Euphorbiaceae) is related to flowering synchrony and tree size, including plant height and circumference at breast height. The study was carried out in the interior and in edges of clearings for gas pipelines and electric lines of a lowland rainforest in south‐eastern Brazil. Monthly observations were performed during one reproductive season, of 19 individuals that grew at edges of electric lines and gas pipelines and at forest interior. Reproductive success was significantly higher at forest interior than at gas pipeline area; there was no significant difference between gas pipeline and electric line areas or between forest interior and electric line area. In the forest edges, only plant height was positively related to plant reproductive success. This is probably related to crown exposure to sunlight, which enhances flower production. At forest interior, reproductive success was positively influenced by the synchrony of flowering activity among neighbouring individuals. In contrast, flowering synchrony based on phenophase intensity negatively impacted reproductive success. Senefeldera verticillata shows temporal dioecy and is mainly pollinated by small social bees, and the high degree of flowering synchrony at low intensity may increase the number of mating partners and therefore enhance its reproductive success. Inside the forest fragments, individuals with thicker trunks showed lower reproductive success, which may be related to a loss of reproductive capacity of older individuals. Our results evidenced the complexity of responses experienced by tropical plants subjected to forest fragmentation because of linear clearings.     

Evolutionary consequences of habitat fragmentation: population size and density affect selection on inflorescence size in a perennial herb

Habitat fragmentation is considered to be one of the major threats to biological diversity worldwide. To date, however, its consequences have mainly been studied in an ecological context, while little is known about its effects on evolutionary processes. In this study we examined whether habitat fragmentation affects selection on plant phenotypic traits via changes in plant-pollinator interactions, using the self-incompatible perennial herb Phyteuma spicatum. Specifically, we hypothesized that limited pollination service in small or low-density populations leads to increased selection for traits that attract pollinators. We recorded mean seed production per capsule and per plant as a measure of pollination intensity and assessed selection gradients (i.e., trait-fitness relationships) in 16 natural populations of varying size and density over 2 years. Mean seed production was not related to population size or density, except for a marginal significant effect of density on the mean number of seeds per capsule in 1 year. Linear selection for flowering time and synchrony was consistent across populations; relative fitness was higher in earlier flowering plants and in plants flowering synchronously with others. Selection on inflorescence size, however, varied among populations, and linear selection gradients for inflorescence size were negatively related to plant population size and density in 1 year. Selection for increased inflorescence size decreased with increasing population size and density. Contrary to our expectation this appeared not to be related to changes in pollination intensity (mean seed production was not related to population size or density in this year), but was rather likely linked to differences in some other component of the abiotic or biotic environment. In summary, our results show that habitat fragmentation may influence selection on plant phenotypic traits, thereby highlighting potential evolutionary consequences of human-induced environmental change.     

The strength of assortative mating for flowering date and its basis in individual variation in flowering schedule

Although it has been widely asserted that plants mate assortatively by flowering time, there is virtually no published information on the strength or causes of phenological assortment in natural populations. When strong, assortative mating can accelerate the evolution of plant reproductive phenology through its inflationary effect on genetic variance. We estimated potential assortative mating for flowering date in 31 old‐field species in Ontario, Canada. For each species, we constructed a matrix of pairwise mating probabilities from the individual flowering schedules, that is the number of flower deployed on successive dates. The matrix was used to estimate the phenotypic correlation between mates, ρ, for flowering date. We also developed a measure of flowering synchrony within species, S, based upon the eigenstructure of the mating matrix. The mean correlation between pollen recipients and potential donors for flowering date was  = 0.31 (range: 0.05–0.63). A strong potential for assortative mating was found among species with high variance in flowering date, flowering schedules of short duration and skew towards early flower deployment. Flowering synchrony, S, was negatively correlated with potential assortment (r = ?0.49), but we go on to show that although low synchrony is a necessary condition for phenological assortative mating, it may not be sufficient to induce assortment for a given phenological trait. The potential correlation between mates showed no seasonal trend; thus, as climate change imposes selection on phenology through longer growing seasons, spring‐flowering species are no more likely to experience an accelerated evolutionary response than summer species.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

Patterns of synchrony in natterjack toad breeding activity and reproductive success at local and regional scales

Empirical studies of the spatiotemporal dynamics of populations are required to better understand natural fluctuations in abundance and reproductive success, and to better target conservation and monitoring programmes. In particular, spatial synchrony in amphibian populations remains little studied. We used data from a comprehensive three year study of natterjack toad Bufo calamita populations breeding at 36 ponds to assess whether there was spatial synchrony in the toad breeding activity (start and length of breeding season, total number of egg strings) and reproductive success (premetamorphic survival and production of metamorphs). We defined a novel approach to assess the importance of short‐term synchrony at both local and regional scales. The approach employs similarity indices and quantifies the interaction between the temporal and spatial components of populations using mixed effects models. There was no synchrony in the toad breeding activity and reproductive success at the local scale, suggesting that populations function as individual clusters independent of each other. Regional synchrony was apparent in the commencement and duration of the breeding season and in the number of egg strings laid (indicative of female population size). Regional synchrony in both rainfall and temperature are likely to explain the patterns observed (e.g. Moran effect). There was no evidence supporting regional synchrony in reproductive success, most likely due to spatial variability in the environmental conditions at the breeding ponds, and to differences in local population fitness (e.g. fecundity). The small scale asynchronous dynamics and regional synchronous dynamics in the number of breeding females indicate that it is best to monitor several populations within a subset of regions. Importantly, variations in the toad breeding activity and reproductive success are not synchronous, and it is thus important to consider them both when assessing the conservation status of pond‐breeding amphibians.     

Genetic structure and AFLP variation of remnant populations in the rare plant Pedicularis palustris (Scrophulariaceae) and its relation to population size and reproductive components

We investigated plant reproduction in relation to genetic structure, population size, and habitat quality in 13 populations of the rare biennial plant Pedicularis palustris with 3-28500 flowering individuals. We used AFLP (amplified fragment length polymorphism) profiles to analyze genetic similarities among 129 individuals (3-15 per population). In a cluster analysis of genetic similarities most individuals (67%) were arranged in population-specific clusters. Analysis of molecular variance indicated significant genetic differentiation among populations and among and within subpopulations (P < 0.001). Gene flow (N(e) m) was low (0.298). On average, plants produced 55 capsules, 17 seeds per fruit, and 42 seedlings in the following growing season. The number of seeds per capsule was independent of population size and of genetic variability. In contrast, the number of capsules per plant (P < 0.05) and the number of seedlings per plant (P < 0.05) were positively correlated with population size. The relation between population size and the number of seeds per plant was not significant (P = 0.075). The number of capsules and of seeds and seedlings per plant (P < 0.01) were positively correlated with genetic variability. Genetic variability was independent of actual population size, suggesting that historical population processes have to be taken into account, too. Stepwise multiple regressions revealed additional significant relationships of habitat parameters (soil pH, C:N ratio), vegetation composition, and standing crop on reproductive components. We conclude that populations of P. palustris are genetically isolated and that reproductive success most likely is influenced by population size, genetic variability, and habitat quality. Management strategies such as moderate grazing, mowing, and artificial gene flow should endeavor to increase population size as well as genetic variation.     

Relationship between population size, allozyme variation, and plant performance in the narrow endemic Cochlearia bavarica

We studied population viability in relation topopulation size and allelic variation in thenarrowly-endemic, monocarpic perennial plantCochlearia bavarica in Bavaria. In 1996,we analysed allelic variation by allozymeelectrophoresis in 24 populations ranging from8–2000 flowering individuals. Fitness-relatedcharacters were investigated in 22 of the 24populations in the field in 1996 (reproductiveand vegetative traits) and 1998 (reproductivetraits only). Differences in allozyme patternwere large between a south-eastern and awestern population group. Genetic diversity,assessed by the Shannon-Wiener diversity index,was low within but high among populations.Small populations had fewer alleles per locus,fewer polymorphic loci, lower observedheterozygosity, and lower genetic diversitythan large populations. Environmentalvariables were not significantly correlatedwith population size or fitness with theexception of light availability, indicatingthat habitat quality was similar for large andsmall populations. Population size showedpositive correlations with number of flowers,fruit set per plant, number of seeds per fruit,and total seed output per plant. Fruit set andnumber of seeds per fruit were positivelycorrelated with the observed heterozygosity andthe proportion of polymorphic loci. We usedpath analyses to study the possible causalrelationships among population size, allelicvariation, and reproductive characters. Thesemodels showed that allelic variation had nodirect influence on reproductive characters,whereas population size did. We conclude thatat present population size reduces viabilityand also reduces allelic variation; but thereduced allelic variation may in the longerterm have negative feed-backs on bothpopulation size and viability.     

Variation of nectar production in relation to plant characteristics in protandrous Aconitum gymnandrum

Aims Floral nectar plays a vital role in plant reproductive success by attracting pollinators. Nectar traits of a flower can depend directly on plant characteristics other than environmental factors and exhibit extensive flower- and plant-level variations. Studies on nectar traits frequently focused on intraplant variation for dichogamous plants, but few have paid attention to both intra- and interplant nectar variations in relation to plant characteristics. Revealing within- and among-plant variation and its relative magnitude is important for our understanding of how pollinator-mediated selection can act on nectar traits and evolution of nectar traits.Methods Through investigating protandrous Aconitum gymnandrum populations at the Alpine Meadows and Wetland Ecosystems Research Station of Lanzhou University, we examined the relationships between nectar production per flower and plant characteristics (e.g. flower position within inflorescences, floral sexual phases, flowering time, inflorescence size and floral attractive traits).Important findings A. gymnandrum exhibited a declining gradient in the nectar volume along inflorescences, with more nectar in basal flowers than distal ones. Protandrous flowers of A. gymnandrum did not show gender-biased nectar production while the nectar volume varied with different stages of floral sexual phases. The significant correlation between the first flowering date of individuals and the mean nectar volume per flower was positive in 2013, but became negative in 2014, suggesting complex effects of biotic and abiotic factors. The mean nectar volume per flower was not related to inflorescence size (the number of total flowers per plant). Furthermore, nectar production was weakly associated with floral attractive traits (the petal width and the galea height), even if the effect of flowering time of individuals was removed, suggesting that the honesty of floral traits as signals of nectar reward for pollinators is not stable in this species.