Correction to: The floral biology and the role of staminal connective appendages during pollination of the endoparasite Bdallophytum americanum (Cytinaceae)

Bdallophytum americanum (Cytinaceae) is an endoparasitic plant species, meaning only the flowers emerge from the host during the reproductive season. Reports on the pollination biology of this species state that its primary pollinators are carrion flies attracted by the smell of the flowers and nectar as a reward. However, the functional role of one of the most outstanding attributes of B. americanum has been neglected. These are the staminal appendages formed by the apical overgrowth of connective tissue during anther development. To determine whether these staminal appendages play a role in pollination, we monitored a nectarless population of B. americanum. We described the inflorescence emergence, floral movements, and pollination and performed field experiments to test whether the absence of the staminal connective appendages affected the visitation frequency. Male inflorescences emerge early, and both male and female flowers open during the day and do not close. Hoverflies are the most frequent visitors to both floral sexes and carry the most pollen. Moreover, the movement of staminal appendages matching the pollen viability changes is reported for the first time. The staminal appendages are the structures where pollinators land before foraging. The field experiments showed that the visitation frequency decreased sharply without staminal appendages. As a landing platform, the staminal connective appendages in B. americanum are crucial for pollinator positioning and collecting viable pollen.

     

Variation in floral longevity in the genus Leptosiphon: mating system consequences

Pollination or fertilisation trigger floral senescence in a wide range of flowering plants, and yet little attention has been given to the implications of this phenomenon to mating system evolution. We examined the effects of pollination on floral senescence in the genus Leptosiphon. Species in the genus exhibit a wide range of breeding systems. In all cases, compatible pollination induced senescence; emasculated flowers lived longer than hand‐outcrossed flowers. In the self‐compatible species, Leptosiphon acicularis and L. bicolor, and in one highly selfing population of L. jepsonii, unmanipulated flowers had reduced longevity compared to emasculated flowers, suggesting that autonomous self‐pollination limits floral longevity in these species. Limited floral longevity in these highly selfing taxa may reduce opportunities for male outcross success, representing a possible source of selection on the mating system. In turn, the mating system might influence how selection acts on floral longevity; obligately outcrossing taxa are expected to benefit from longer floral longevities to maximise opportunities for pollination, while selfing taxa might benefit from earlier floral senescence to reduce resource expenditure. Overall, the longevity of unpollinated flowers increased with the level of outcrossing in the genus Leptosiphon. Our results taken together with those of a previous study and similar results in other species suggest that floral longevity may represent a largely unexamined role in mating system evolution.     

The influence of nectar production and floral visitors on the female reproductive success of Inga (Fabaceae): a field experiment

Floral morphology, nectar secretion strategies and the contribution of pollinators to the reproductive success of plants provide important clues regarding the levels of generalization or specialization in pollination systems. Anthesis throughout the day and night allows flowers to be visited by diurnal and nocturnal pollinators, promoting generalization or specialization. We studied three species in the diverse tropical genus Inga to: (1) quantify the response of flowers to successive nectar extractions and (2) determine the contribution of diurnal and nocturnal floral visitors to female reproductive success. Inga flowers could be clearly distinguished mainly on the basis of the staminal tube diameter and the quantities of filaments and pollen grains. Successive nectar removals led to a decrease of 60% in the total nectar secretion in I. vera and to increases of 20% in I. ingoides and 10% in I. striata. Despite these differences, the studied Inga spp. exhibited similar patterns of visitation rates and shared diurnal and nocturnal pollinators. Nocturnal pollinators contributed ten times more than diurnal pollinators to the female reproductive success of Inga. Floral morphology, nectar secretion patterns and pollination ecology data suggest an evolutionary trend towards specialization for nocturnal pollinators in Inga spp. with crepuscular or nocturnal flowers. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 230–245.     

Relative impact of mate versus pollinator availability on pollen limitation and outcrossing rates in a mass‐flowering species

Plant mating systems are driven by several pre‐pollination factors, including pollinator availability, mate availability and reproductive traits. We investigated the relative contributions of these factors to pollination and to realized outcrossing rates in the patchily distributed mass‐flowering shrub Rhododendron ferrugineum. We jointly monitored pollen limitation (comparing seed set from intact and pollen‐supplemented flowers), reproductive traits (herkogamy, flower size and autofertility) and mating patterns (progeny array analysis) in 28 natural patches varying in the level of pollinator availability (flower visitation rates) and of mate availability (patch floral display estimated as the total number of inflorescences per patch). Our results showed that patch floral display was the strongest determinant of pollination and of the realized outcrossing rates in this mass‐flowering species. We found an increase in pollen limitation and in outcrossing rates with increasing patch floral display. Reproductive traits were not significantly related to patch floral display, while autofertility was negatively correlated to outcrossing rates. These findings suggest that mate limitation, arising from high flower visitation rates in small plant patches, resulted in low pollen limitation and high selfing rates, while pollinator limitation, arising from low flower visitation rates in large plant patches, resulted in higher pollen limitation and outcrossing rates. Pollinator‐mediated selfing and geitonogamy likely alleviates pollen limitation in the case of reduced mate availability, while reduced pollinator availability (intraspecific competition for pollinator services) may result in the maintenance of high outcrossing rates despite reduced seed production.     

Floral traits and mating system of Hibiscus trionum (Malvaceae)

Variations in floral traits and floral structures influence plant mating systems. Hibiscus trionum produces large, showy flowers typical of an outcrossing species, yet flowers are autonomously self-pollinated. In this study, we measured floral morphology, breeding system and outcrossing rate estimated by ISSR markers. Results indicate that two types of flowers were observed in H. trionum, and the type I with bigger petals appears to be much more visible to pollinators, demonstrated by than type II flowers with smaller petals. The flowers with hand pollination were closed 1 h earlier than intact flowers, whether they were type I or II. The relationship between the amount of pollen deposited on the stigma and the number of seeds per capsule was highly significant, and 80 or more pollens per flower can make the mean number of seeds (mean = 37) in H. trionum. Delayed selfing in H. trionum did not provide a large contribution to seed production, since reproductive assurance were only 0.025. However, successful reproduction of 72.5% flowers in the absence of pollinators suggested that selfing provides reproductive assurance during seasons, in which pollinators were limiting. The multilocus outcrossing rates in different populations varied from 0.982 to 1.200, with a mean of 1.116. Our data provide an empirical demonstration of a predominantly outcrossing species with potential delayed selfing when pollinators are absent or scarce.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE. XI. THE FLORAL BIOLOGY OF NELUMBO PENTAPETALA

The floral biology of Nelumbo pentapetala (Walter) Fernald, the American lotus, native to Texas, was investigated. Anthesis occurs over three consecutive days with flowers opening each morning and closing around noon. First-day flowers are protogynous with the perianth parts partially expanded so that pollen-covered insects which are attracted by floral color and the intense “fruity” odor (diffused with the aid of increased floral temperature) are directed on to the flattened receptacle (= carpellary receptacle) from which the receptive stigmas protrude, thus accomplishing pollination. During the second morning anther dehiscence begins and insects which visit and forage within the flower become covered with pollen and typically crawl over the still receptive stigmas achieving “facilitated” self-pollination (indirect autogamy). By mid-morning of the second day the stigmas dry and become non-receptive to pollen. During the third day of anthesis perianth and staminal parts quickly abscise and over the period of a few weeks the receptacle and enclosed fruits mature. In most populations studied, Hymenoptera (e.g., Lusioglossum spp., and Apis mellifera) were the most abundant and effective pollinators. In some populations, however, Coleoptera (e.g., Chauliognathus) were also numerous and effective pollinators. It is suggested that the overall floral structure (e.g., large numbers of stamens, masses of pollen, staminal appendages) are adaptations which facilitate the pollination of Nelumbo by beetles.     

Reproductive correlates of mating system variation in Eichhornia paniculata (Spreng.) Solms (Pontederiaceae)

Comparative studies of related plant species indicate that evolutionary shifts in mating systems are accompanied by changes in reproductive attributes such as flower size, floral morphology, and pollen/ovule ratio. Recent theoretical work suggests that patterns of investment in reproduction should also change with the mating system. In a glasshouse study, we investigated the extent to which mating system differences among populations of Eichhornia paniculata (Pontederiaceae) were correlated with changes in allocation to male and female function, floral display, and the regulation of investment in reproduction through fruit and ovule abortion. Significant differences in the amount of biomass allocated to reproductive structures were evident among six populations of E. paniculata. As predicted by sex allocation theory, the proportion of dry weight allocated to male function decreased with the outcrossing rate of populations. Six of the eight attributes used to characterize floral display also differed significantly among populations. However, with the exception of two attributes describing the number of flowers produced by inflorescences, these were not correlated with outcrossing rate. Levels of fruit and ovule abortion were determined in two populations with contrasting mating systems under different nutrient and pollination treatments. Virtually all fruits initiated by plants from a self-fertilizing population were matured, while the amount of fruit abortion in an outcrossing population increased with flower production. Ovule abortion was low in both populations. Our results demonstrate that the evolution of self-fertilization in E. paniculata is associated with changes in investment to reproduction that normally distinguish selfing and outcrossing species.     

Floral longevity and autonomous selfing are altered by pollination and water availability in Collinsia heterophylla

Background and Aims

A plant investing in reproduction partitions resources between flowering and seed production. Under resource limitation, altered allocations may result in floral trait variations, leading to compromised fecundity. Floral longevity and timing of selfing are often the traits most likely to be affected. The duration of corolla retention determines whether fecundity results from outcrossing or by delayed selfing-mediated reproductive assurance. In this study, the role of pollination schedules and soil water availability on floral longevity and seed production is tested in Collinsia heterophylla (Plantaginaceae).

Methods

Using three different watering regimes and pollination schedules, effects on floral longevity and seed production were studied in this protandrous, flowering annual.

Key Results

The results reveal that soil water status and pollination together influence floral longevity with low soil water and hand-pollinations early in the floral lifespan reducing longevity. However, early pollinations under excess water did not extend longevity, implying that resource surplus does not lengthen the outcrossing period. The results also indicate that pollen receipt, a reliable cue for fecundity, accelerates flower drop. Early corolla abscission under drought stress could potentially exacerbate sexual conflict in this protandrous, hermaphroditic species by ensuring self-pollen paternity and enabling male control of floral longevity. While pollination schedules did not affect fecundity, water stress reduced per-capita seed numbers. Unmanipulated flowers underwent delayed autonomous selfing, producing very few seeds, suggesting that inbreeding depression may limit benefits of selfing.

Conclusions

In plants where herkogamy and dichogamy facilitate outcrossing, floral longevity determines reproductive success and mating system. Reduction in longevity under drought suggests a strong environmental effect that could potentially alter the preferred breeding mode in this mixed-mated species. Extrapolating the findings to unpredictable global drought cycles, it is suggested that in addition to reducing yield, water stress may influence the evolutionary trajectory of plant mating system.     

Pollinator directionality as a response to nectar gradient: promoting outcrossing while avoiding geitonogamy

Plants with multiple flowers could be prone to autonomous self‐pollination and insect‐mediated geitonogamy, but physiological and ecological features have evolved preventing costs related to autogamy. We studied the rare perennial herb Dictamnus albus as a model plant, with the aim of describing the plant–pollinator system from both plant and pollinator perspectives and analysing features that promote outcrossing in an entomophilous species. The breeding system and reproductive success of D. albus were investigated in experimental and natural conditions, showing that it is potentially self‐compatible, but only intra‐inflorescence insect‐mediated selfing is possible. Nectar analysis showed gender‐biased production towards the female phase, which follows the male phase, and during flowering, full blooming is found in flowers at the bottom of the raceme. Among a wide spectrum of insect visitors, three genera (Bombus, Apis, Megachile) were found to be principal pollinators. A study of insect behaviour showed a tendency towards bottom‐to‐top flights for the most important pollinators Bombus spp. and Apis mellifera: upward movements on the racemes could be explained by foraging behaviour, from more to less rewarding flowers. In accordance with the ‘declining reward hypothesis’, bumblebees and honeybees leave the plant when gain of reward is low, after which few flowers are visited, reducing the chance of self‐pollen transfer among flowers. Intra‐flower self‐pollination is prevented in D. albus by protandry and herkogamy, while the nectar‐induced sequential pattern of pollinator visits avoids geitonogamy and tends to maximise pollen export, promoting outcrossing. All these features for preventing selfing benefit plant fitness and population genetic structure.     

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.     

BREEDING SYSTEM EVOLUTION IN LEAVENWORTHIA: BREEDING SYSTEM VARIATION AND REPRODUCTIVE SUCCESS IN NATURAL POPULATIONS OF LEAVENWORTHIA CRASSA (CRUCIFERAE)

Populations of Leavenworthia crassa (Cruciferae) studied for 3 years exhibited among- and within-population genetic variation for a suite of floral and reproductive traits (flower width, petal length, anther position, ability to set seeds in the absence of pollinators, time to first flowering) associated with breeding system. We used electrophoretic markers to show that a population with small, monomorphically colored flowers with introrse anthers had a significantly lower outcrossing rate (t = 0.03) than did a population with larger, polymorphically colored flowers with extrorse anthers (t = 0.33). In the more-outcrossing population the correlation between higher maternal plant outcrossing rate and the suite of six traits approached significance (P < 0.067), with greater petal size, greater flower width, and reduced ability to set seeds in the absence of pollinators contributing significantly. Plants in selfing populations had a generally higher reproductive success, with a higher number of flowers per plant, a smaller proportion of unfertilized ovules, a smaller proportion of fertilized ovules aborted, a higher rate of fruit set, and overall a larger number of seeds matured than did plants from the more outcrossing populations. Pollen limitation did not appear to account for lower reproductive success in outcrossing populations. Resource limitation did not differ substantially between populations. However, within-ovary patterns of fertilization, abortion, and seed weight were significantly less random in outcrossing populations than in selfing populations, suggesting that differential gamete and embryo success may be responsible for lower reproductive success in outcrossing populations.     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads

Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self‐compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self‐compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.     

Day–night fluctuations in floral scent and their effects on reproductive success in <Emphasis Type="Italic">Lilium auratum</Emphasis>

We examined the contribution of diurnal and nocturnal pollination to male and female reproductive success in Lilium auratum. Plants were bagged for either 12 h during the day or at night to allow either only nocturnal or only diurnal visitors to forage throughout the flowering period. We found that there was no significant difference in the seed:ovule ratio among diurnally pollinated, nocturnally pollinated, or control flowers. However, in terms of male reproductive success, it was more advantageous for the plants to be pollinated both diurnally and nocturnally: the numbers of pollen grains remaining in diurnally pollinated or nocturnally pollinated flowers were significantly greater than those in control flowers. The total amount of floral volatiles of L. auratum was significantly higher at night than during the day. The constituents of floral scent of all time series examined were mostly monoterpenoids, many of which serve as attractants for nocturnal hawkmoths. Such nocturnally biased floral scent emission of L. auratum might achieve male reproductive success by attracting nocturnal visitors, which may suggest that the relative contribution of floral scent in this species is biased towards male reproductive success.     

Floral biology of Eriope blanchetii (Lamiaceae) in coastal sand dunes of NE Brazil

Abstract The floral biology of the sand dune shrub Eriope blanchetii (Benth) Harley was investigated in a wild population located within an environment protection area near Salvador, NE Brazil. Inflorescence and flower development were monitored, and the reproductive biology was assessed. Inflorescences are terminal panicles, each bearing from one to 38 small, zygomorphic flowers. The duration of an inflorescence is about 2 months, with few new flowers each day. The rewards to visitors are pollen and nectar, the latter being produced in small quantities at the corolla base, near the stylopodium. Three sequential phases of flower development were identified, based on colour changes, pollen and nectar availability, and stigmatic receptivity. Anthesis takes place throughout the day within each plant, and flowers are strongly protandrous. Flowers are short‐lived, with individual variations of 7–30 h until corolla abscission. This variation is probably associated to a post‐pollinating response, which reduces the length of the female phase and could be advantageous for the economy of nectar production and for increased efficient pollination of the remaining flowers on a plant. The results of the pollination tests (apomixis, spontaneous and manual autogamy, manual cross‐pollination, and control) reinforced the role of pollinators for the reproductive success of E. blanchetii, given that the production of fruits from spontaneous pollination is unlikely to be resulting from protandry and given the morphological floral features. Taken together, the morphological and physiological aspects of E. blanchetii floral biology favour cross‐pollination, and it is likely that most of the fruits in nature are produced by outcrossing.     

Pollination ecology of Isoglossa woodii,a long‐lived,synchronously monocarpic herb from coastal forests in South Africa

Synchronous monocarpy in long‐lived plants is often associated with pollination by wind, in part because infrequent mass flowering may satiate pollinators. Selfing in synchronous monocarps may provide reproductive assurance but conflict with the benefits of outcrossing, a key evolutionary driver of synchrony. We predicted that animal‐pollinated species with synchronous flowering would have unspecialised flowers and attract abundant generalised pollinators, but predictions for selfing and outcrossing frequencies were not obvious. We examined the pollination biology of Isoglossa woodii (Acanthaceae), an insect‐pollinated, monocarpic herb that flowers synchronously at 4–7‐year intervals. The most frequent visitor to I. woodii flowers was the African honeybee, Apis mellifera adansonii. Hand‐pollination failed to enhance seed production, indicating that the pollinators were not saturated. No seed was set in the absence of pollinators. Seed set was similar among selfed and outcrossed flowers, demonstrating a geitonogamous mixed‐mating strategy with no direct evidence of preferential outcrossing. Flowers contained four ovules, but most fruits only developed one seed, raising the possibility that preferential outcrossing occurs by post‐pollination processes. We argue that a number of the theoretical concerns about geitonogamous selfing as a form of reproductive assurance do not apply to a long‐lived synchronous monocarp such as I. woodii.     

片断化生境中濒危植物黄梅秤锤树的开花生物学、繁育系统与生殖成功的因素

 研究处于片断化生境中濒危植物的繁育系统、传粉过程及生殖成功状况, 可为评估物种的受胁迫程度及制订相应的保护和管理策略提 供重要依据。通过野外观察和人工授粉实验等方法,对片断化生境中濒危植物黄梅秤锤树(Sinojackia huangmeiensis)的开花生物学和繁育系统 进行了初步研究, 并探讨了影响其生殖成功的因素。黄梅秤锤树的总状花序上通常有两性花2～6朵, 单花花期约5～7 d, 单花花药不同步开裂 使整朵花花粉活力持续时间延长。虽然花开放过程中柱头始终高于花药, 在单花水平上可在一定程度上避免同花自交的发生, 但由于花粉活力 和柱头可授期之间有一定的重叠期, 在自然情况下, 由于外界媒介动物和风媒的作用, 不可避免地会产生同株同花和同株异花传粉。实验结果 表明, 该种不存在无融合生殖现象, 繁育类型以异交为主, 部分自交亲和且需要传粉者。主要的传粉昆虫为黑带食蚜蝇(Epistrophe balteata) 、中华蜜蜂 (Apis cerana) 和中华回条蜂(Habropoda sinensis)。黄梅秤锤树自然状态下的结实率较低(10.33%), 平均每果种子数较少(0.387 个), 这主要与传粉昆虫少、花粉传递效率低, 以及柱头上自花及同株异花花粉的落置和自交亲和性低有关。但是单株较大的开花量保证了一定 的果实和种子产量, 使居群能自然更新。部分自交亲和性可能为该物种适应生境片断化提供了一定的生殖保障及进化潜力。对该残存居群的保 护, 应着眼于加强对传粉昆虫赖以生存的自然生态系统的恢复。     

Coexistence of Delayed Autonomous Self pollination and Deceptive Pollination in Arthropodium cirratum (Asparagaceae)

Delayed autonomous pollination and deceptive pollination are assumed to be relatively common in flowering plants, but no species have been reported to use both of these cunning reproductive strategies. In this study, we examined whether delayed selfing and mimicry were used concurrently in Arthropodium cirratum. Flowers of A.cirratum were manipulated to assess whether their stamen appendages and corolla closing movements were functional in deceptive pollination and delayed selfing, respectively. Our results indicated that anther or pollen imitation of the yellow stamen appendages contributed to 93 per cent of successful mimetic attraction, and this mimicry was an important driver of cross pollination of the species. In addition, we observed closure of the perianth relocated the stamens over the stigma at the end of anthesis, which significantly increased the average seed number per fruit of intact flowers over flowers emasculated before closure (20.62 versus 11.79). Our findings confirmed the coexistence of delayed autonomous self pollination and deceptive pollination in A.cirratum. Our results also suggested that delayed selfing could add fitness benefits for this mimetic attraction species.     

新西兰岩石百合中两种共存的繁育策略：延迟自动自交和欺骗传粉吸引

延迟自动自交和欺骗传粉吸引在被子植物多数类群中有相关报道,但是至今没有发现两种繁育策略在同一物种中共存现象。本研究通过对照试验检测新西兰岩石百合雄蕊附属物和花冠闭合运动是否分别具有欺骗吸引和延迟自交功能。研究结果表明,新西兰岩石百合黄色的雄蕊附属物拟态花粉（或花药）,约93%的昆虫访花行为源自黄色花药附属物的欺骗吸引,雄蕊附属物的报酬拟态功能有效提高昆虫拜访频率并促进异交。同时,研究发现新西兰岩石百合花期结束时花瓣闭合促使雌雄隔离距离的缩小,花瓣自然闭合的花朵平均结籽数（20.62）显著高于闭合前去雄处理花朵（11.79）。我们的结果表明延迟自动自交与欺骗传粉吸引两种繁育策略在新西兰岩石百合中共存。     

Flexible pollination system in an unpalatable shrub Daphne miyabeana (Thymelaeaceae)

The stability of plant reproductive success is dependent on the reproductive system of the plant and the surrounding pollinator assemblages. Here we documented the pollination ecology of a shrub, Daphne miyabeana (Thymelaeaceae), that is unpalatable to deer in two different habitats (open and closed canopy) at a site with severe deer browsing. We found that 31.22 ± 3.07% (mean ± SE) and 9.01 ± 1.71% of the total number of D. miyabeana flowers become fruits in open and closed habitats, respectively. Although spontaneous selfing was possible, the fruit set of that was much lower than the fruit set of an artificial outcrossing treatment. In the closed habitat, the fruit set of the natural pollination was lower than that of artificial outcrossing, whereas in the open habitat the fruit set of those two were similar. In the open habitat, small bees were the major flower visitors, whereas in the closed habitat, ants were the major flower visitors, which did not contribute to pollination. Our findings suggest that D. miyabeana is pollinated by occasional visits of small bees, but it is also self‐compatible and spontaneous selfing is possible. Although its pollination is shaped by abiotic environmental factors, D. miyabeana can reproduce in different habitat conditions. These flexible reproductive systems may be an advantage in coping with the recent environmental change in forest habitats.