Pollination and floral scent differentiation in species of the Philodendron bipinnatifidum complex (Araceae)

The Philodendron bipinnatifidum complex of Philodendron subgenus Meconostigma may comprise four species, which because of only slight and not very distinct morphological differences are not all unanimously recognized as good species. To find out whether these species are reproductively isolated, we studied the flowering rhythm, thermogenesis and pollination biology of three species of this complex, namely of P. bipinnatifidum, P. aff. bipinnatifidum (provisionally named “P. form selloum”) and P. mello-barretoanum in Brazil. Of the first two mentioned taxa, floral scent was collected and scent compounds were identified by GC–MS. The results showed that the coastal forest species P. bipinnatifidum has a two-, or three-night flowering rhythm, with the pistillate stage in the first night and the staminate stage lasting the second and sometimes also the third night. Strong thermogenesis with extended heating periods of several hours during the first part of the usual two subsequent nights and the maximum temperatures of up to 40 °C absolute heating of the spadices occurred in the pistillate and staminate stages. Concomitant with the heating periods, relatively low amounts of principally (Z)-2-pentenyl acetate and (Z)-jasmone were emitted by both the pistillate and staminate stage inflorescences. The dynastid scarab beetle Cyclocephala variolosa was the only pollinator attracted. The upland forest P. form selloum always had a two-night flowering rhythm with the pistillate stage in the first and the staminate stage in the subsequent night. This world-record holder of thermogenesis can heat up to the remarkable 45 °C during a relatively short period in the evening of the pistillate stage. During the thermogenic period, enormous amounts of principally 4-methoxystyrene and 3,4-dimethoxystyrene were produced and which could attract a large number of female and male individuals of the dynastid scarab beetle Erioscelis emarginata. In the staminate stage of P. form selloum, temperature elevation is significantly lower and the scent compounds are different from the pistillate stage. The cerrado biome species P. mello-barretoanum has a flowering rhythm similar to P. form selloum, reaching a maximum heating of about 40 °C during the pistillate stage. The sole pollinator attracted was Cyclocephala atricapilla. The differences observed and analyzed among the taxa, including the flowering rhythm, thermogenic activities, scent compounds emitted, pollinating dynastid scarab beetles attracted, as well as slight morphological differences and apparent geographical exclusiveness noted in these three taxa are strong indicators that P. bipinnatifidum, P. form selloum and P. mello-barretoanum are different enough to be considered good species. The morphological affinities of these species might be a hint that speciation has been a recent event and/or also that reproductive isolation based on different, non-overlapping distribution areas, different scent compounds and different pollinators was effective enough to need further morphological differentiation.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Floral volatiles fromClarkia breweri andC. concinna (Onagraceae): Recent evolution of floral scent and moth pollination

Clarkia breweri (Onagraceae) is the only species known in its genus to produce strong floral fragrance and to be pollinated by moths. We used gas chromatography-mass spectrometry (GC-MS) to identify 12 abundant compounds in the floral headspace from two inbred lines ofC. breweri. These volatiles are derived from two biochemical pathways, one producing acyclic monoterpenes and their oxides, the other leading from phenylalanine to benzoate and its derivatives. Linalool and linalool oxide (pyran form) were the most abundant monoterpenoids, while linalool oxide (furan form) was present at lower concentrations. Of the aromatic compounds detected, benzyl acetate was most abundant, whereas benzyl benzoate, eugenol, methyl salicylate, and vanillin were present as minor constituents in all floral samples. The two inbredC. breweri lines differed for the presence of the additional benzenoid compounds isoeugenol, methyleugenol, methylisoeugenol, and veratraldehyde. We also analyzed floral headspace fromC. concinna, the likely progenitor ofC. breweri, whose flowers are odorless to the human nose. Ten volatiles (mostly terpenoids) were detected at low concentrations, but only when headspace was collected from 20 or more flowers at a time. Trans--ocimene was the most abundant floral compound identified from this species. Our data are consistent with the hypothesized recent evolution of floral scent production and moth pollination inC. breweri.     

Pollination system of Corylopsis gotoana (Hamamelidaceae) and its stonefly (Plecoptera) co‐pollinator

The order Plecoptera is primarily known for its aquatic nymphs and their importance in many stream ecosystems. However, the biology and life history of Plecopteran terrestrial adults is mostly unknown, and hence interactions with plants have rarely been documented. Here, we report on insect visitation to flowers of Corylopsis gotoana (Hamamelidaceae) in a temperate forest of Japan, with emphasis on the behavior and pollination role of adults of a stonefly species, Strophopteryx nohirae (Taeniopterygidae), which were frequently observed in the flowers. The most frequent insect visitors to Corylopsis flowers were Bombylius major (Bombyliidae), Apis cerana (Apidae) and Bombus ardens (Apidae), all of which were considered main pollinators because of high rates of visitation and pollen attachment. Strophopteryx nohirae frequently visited Corylopsis trees, on which they foraged for pollen and mated. A field experiment on the pollination success of Corylopsis flowers visited by S. nohirae verified that this stonefly contributed to pollination.     

Phenotypic selection on floral traits in the arctic plant Parrya nudicaulis (Brassicaceae)

The evolution of floral traits is often attributed to pollinator‐mediated selection; however, the importance of pollinators as selective agents in arctic environments is poorly resolved. In arctic and subarctic regions that are thought to be pollen limited, selection is expected to either favor floral traits that increase pollinator attraction or promote reproductive assurance through selfing. We quantified phenotypic selection on floral traits in two arctic and two subarctic populations of the self‐compatible, but largely pollinator‐dependent, Parrya nudicaulis. Additionally, we measured selection in plants in both open pollination and pollen augmentation treatments to estimate selection imposed by pollinators in one population. Seed production was found to be limited by pollen availability and strong directional selection on flower number was observed. We did not detect consistently greater magnitudes of selection on floral traits in the arctic relative to the subarctic populations. Directional selection for more pigmented flowers in one arctic population was observed, however. In some populations, selection on flower color was found to interact with other traits. We did not detect consistently stronger selection gradients across all traits for plants exposed to pollinator selection relative to those in the pollen augmentation treatment; however, directional selection tended to be higher for some floral traits in open‐pollinated plants.     

Interannual fluctuations in floral longevity, pollinator visitation and pollination limitation of an alpine plant (Gentiana straminea Maxim., Gentianaceae) at two altitudes in the Qinghai-Tibetan Plateau

Prolonged floral longevity and bumblebees as dominate pollinators in alpine ecosystem have been suggested to overcome pollination limitation of alpine plants arising from the decrease of pollinator activity with increasing altitude. However, this conclusion has never been examined in the Qinghai-Tibetan Plateau (QTP), the highest and largest plateau in the world. In this study, we intended to test year-to-year correlations between floral longevity, visiting frequency and pollen limitation of this species between two populations (at 3200 m and 4000 m) of Gentiana straminea in this plateau. Pollinator exclusion elongated both male and female phases greatly at both sites, and durations of both male and female phases in natural condition varied greatly over three years. The visiting frequency of bumblebees varied greatly at the higher altitude, but seemed to be stable at the lower altitude. Seed production was pollination limited in both populations in most studied years. The floral durations, pollinator frequency and pollination limitation showed no significant and consistent variations with the increase of altitude. The previous hypothesis that the prolonged floral longevity of alpine plants can compensate for low levels of pollinator visitation therefore could not be confirmed, and our results further suggested that in the QTP platform, the altitude shows no consistent effect on the reproductive performance of this species, despite that the fluctuation of visit frequency intensified at the higher altitude.     

Pollen:ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): an evaluation with Argentine species

The pollen:ovule ratio (P/O) has traditionally been used as a rough estimator of plant breeding systems. It has been shown that plant breeding systems are associated with particular floral traits. In this study, we determined the P/O in 21 Leguminosae species from Argentina and explored relationships between P/O and taxonomic position, flower size, floral rewards, pollen presentation and pollination mechanisms. According to the results, 15 out of the 21 species classified were obligate xenogamous, although some of them have been recorded as facultative xenogamous in previous studies. There was a significant effect of taxonomic position (genus), reward type and pollination mechanism on P/O. Species offering only nectar as a floral reward (which were species with a brush mechanism) had a significantly lower P/O than species offering pollen or pollen and nectar. Species with the brush pollination mechanism had the lowest P/O, while species with valvular and pump mechanism had the highest P/O. However, pollen presentation (primary and secondary) and flower size did not have a significant effect on P/O. Our results demonstrate that P/O variability is determined by taxonomic position and pollination mechanism in this plant group.     

Breeding systems in Clivia (Amaryllidaceae): late‐acting self‐incompatibility and its functional consequences

Late‐acting (ovarian) self‐incompatibility, characterized by minimal or zero seed production following self‐pollen tube growth to the ovules, is expected to show phylogenetic clustering, but can otherwise be difficult to distinguish from early‐acting inbreeding depression. In Amaryllidaceae, late‐acting self‐incompatibility has been proposed for Narcissus (Narcisseae) and Cyrtanthus (Cyrtantheae). Here, we investigate whether it occurs in the horticulturally important genus Clivia (Haemantheae) and test whether species in this genus experience ovule discounting in wild populations. Seed‐set results following controlled hand pollinations revealed that Clivia miniata and C. gardenii are largely self‐sterile. Self‐ and cross‐pollinated flowers of both species had similar proportions of pollen tubes entering the ovary, and those of C. gardenii also did not differ in the proportions of pollen tubes that penetrated ovules, thus ruling out classical gametophytic self‐incompatibility acting in the style, but not early inbreeding depression. Flowers that received equal mixtures of self‐ and cross‐pollen set fewer seeds than those that received cross‐pollen only, but it was unclear whether this effect was a result of ovule discounting or interactions on the stigma. The prevention of self‐pollination by the emasculation of either single flowers or whole inflorescences in wild populations did not affect seed set, suggesting that ovule discounting is not a major natural limitation on seed production. Flowers typically produce one to three large fleshy seeds from approximately 16 available ovules, even when supplementally hand pollinated, suggesting that fecundity is mostly resource limited. The results of this study suggest that Clivia spp. are largely self‐sterile as a result of either a late‐acting self‐incompatibility system or severe early inbreeding depression, but ovule discounting caused by self‐pollination is not a major constraint on fecundity. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 155–168.     

三室算盘子寄主上弯头细蛾的生物学特性及花气味成分分析

【目的】为了从生物学和化学生态学角度探讨弯头细蛾Epicephala ancylopa和寄主三室算盘子Glochidion sp.间专性传粉的互利共生关系稳定性。【方法】本研究在野外观察和室内实验的基础上,对专性传粉育幼互利共生体系中三室算盘子、弯头细蛾生物学特性进行详细研究,探究互利共生双方利益得失;用动态顶空吸附法分别收集三室算盘子雄花和雌花气味物质,运用气相色谱-质谱联用技术(GC-MS)分析并鉴定其有效成分,用峰面积归一化与内标法定性定量;最后通过主成分分析法比较雄花和雌花之间气味化学成分的差异性。【结果】弯头细蛾在云南省普洱太阳河国家森林公园每年有1个世代,成虫和幼虫的活动时间分别在3-4月和8-10月。三室算盘子结实率为44.20%,被蛀食率为69.94%,平均每头幼虫消耗2.55枚种子来满足自身生长发育,寄主植物留有83.06%完好的种子,以维持互利共生关系的稳定。三室算盘子雌雄花气味中共鉴定出24种挥发物,主要以单萜类和倍半萜类物质为主,其中(Z)-罗勒烯和β-榄香烯两种萜类物质含量最高(分别为47.11%和22.72%),推测其是吸引弯头细蛾传粉的主要气味成分;雄花和雌花之间气味化学成分存在明显的差异,具有两性异型性。【结论】弯头细蛾通过以卵越夏和以蛹越冬对策,实现成虫发生期与三室算盘子花期的精准匹配。弯头细蛾成虫白天静伏,傍晚开始活动,三室算盘子花的气味物质也只在晚上才明显释放,且雌雄花气味化学成分的两性异型性有利于弯头细蛾辨别雌雄花,以完成采集花粉与传粉行为。该研究结果为头细蛾属昆虫与算盘子属植物专性传粉互利共生关系稳定性的维持机制提供了新的依据,也为深入开展通过触角电生理检测和生物行为实验来筛选吸引传粉头细蛾的活性物质提供了理论依据。     

Description of a new species of the genus Resseliella (Diptera: Cecidomyiidae), a pollinator of Kadsura longipedunculata (Schisandraceae) in China,with comments on its flower‐visiting habit

A gall midge pollinating flowers of Kadsura longipedunculata is described as Resseliella kadsurae sp. nov. based on adult specimens collected in 2006 from Langshan, Xinning County, Hunan Province, China. The new species is distinguishable from other known congeners in the combination of the following characteristics: gynecoid male flagellomeres, mottled wings, simple claws on all legs; male hypoproct with a transverse bridge‐like structure ventrally connecting both sides of hypoproct. In particular, the gynecoid male flagellomeres and transverse bridge‐like structure are unique to the new species. Partial sequencing of the cytochrome oxidase subunit I mitochondrial gene indicates that R. kadsurae is grouped with the R. theobardi + R. yagoi clade, but this clade and the monophyly of the genus Resseliella are not statistically supported. The new species is nocturnal, and mating occurs at dusk. At the time of mating, males and females hang on opposite sides of non‐sticky old spider webs in a venter‐to‐venter position. Immature stages and the life history of R. kadsurae are unknown. The flower‐visiting habit of R. kadsurae is contrasted with that of other flower‐visiting gall midges and the probability that this behavior exists in other congeners is discussed.     

Floral volatiles, pollinator sharing and diversification in the fig-wasp mutualism: insights from Ficus natalensis, and its two wasp pollinators (South Africa)

Combining biogeographic, ecological, morphological, molecular and chemical data, we document departure from strict specialization in the fig-pollinating wasp mutualism. We show that the pollinating wasps Elisabethiella stuckenbergi and Elisabethiella socotrensis form a species complex of five lineages in East and Southern Africa. Up to two morphologically distinct lineages were found to co-occur locally in the southern African region. Wasps belonging to a single lineage were frequently the main regional pollinators of several Ficus species. In South Africa, two sister lineages, E. stuckenbergi and E. socotrensis, pollinate Ficus natalensis but only E. stuckenbergi also regularly pollinates Ficus burkei. The two wasp species co-occur in individual trees of F. natalensis throughout KwaZulu-Natal. Floral volatile blends emitted by F. natalensis in KwaZulu-Natal were similar to those emitted by F. burkei and different from those produced by other African Ficus species. The fig odour similarity suggests evolutionary convergence to attract particular wasp species. The observed pattern may result from selection for pollinator sharing among Ficus species. Such a process, with one wasp species regionally pollinating several hosts, but several wasp species pollinating a given Ficus species across its geographical range could play an important role in the evolutionary dynamics of the Ficus-pollinating wasp association.     

黑面神雌花气味传粉前后的变化及其生态意义

该文以与头细蛾传粉相关的叶下珠科植物黑面神为材料,采用动态顶空吸附法分别收集了该物种雌花传粉前后释放的挥发物,利用气相色谱-质谱联用技术分析了其化学成分,探索了三者间花气味的异同,并对其生态意义进行了讨论。结果表明:从该物种传粉前、传粉期和传粉后的雌花气味中共分离出22种挥发物(传粉前12种,传粉期15种和传粉后13种),以脂肪族衍生物和萜类物质为主。3-己烯醛、反-2-己烯醛、顺-3-己烯-1-醇、顺-乙酸-3-己烯酯、反-β-罗勒烯、β-石竹烯、β-蛇麻烯、反,反-α-金合欢烯和长叶烯是其主要挥发物。传粉前后雌花气味的化学种类及组成发生了显著分化,且这种变化主要是由于各时期主要挥发物发生改变而引起的。反-β-罗勒烯在传粉期含量最高,而在传粉后显著下降,3-己烯、反-2-己烯醛、顺-3-己烯-1-醇和顺-乙酸-3-己烯酯在传粉前含量均最高,β-石竹烯含量在传粉后显著升高,且长叶烯仅在传粉后出现。在传粉期和传粉后β-蛇麻烯和反,反-α-金合欢烯的含量均显著上升。同时,传粉期雌花的释放量明显高于传粉前和传粉后的。综上结果显示,黑面神雌花传粉前后花气味发生了质与量的改变。初步推测黑面神雌花...     

Altitudinal flower size variation correlates with local pollinator size in a bumblebee‐pollinated herb,Prunella vulgaris L. (Lamiaceae)

The influence of locally different species interactions on trait evolution is a focus of recent evolutionary studies. However, few studies have demonstrated that geographically different pollinator‐mediated selection influences geographic variation in floral traits, especially across a narrow geographic range. Here, we hypothesized that floral size variation in the Japanese herb Prunella vulgaris L. (Lamiaceae) is affected by geographically different pollinator sizes reflecting different pollinator assemblages. To evaluate this hypothesis, we posed two questions. (1) Is there a positive correlation between floral length and the proboscis length of pollinators (bumblebees) across altitude in a mountain range? (2) Does the flower–pollinator size match influence female and male plant fitness? We found geographic variation in the assemblage of pollinators of P. vulgaris along an altitudinal gradient, and, as a consequence, the mean pollinator proboscis length also changed altitudinally. The floral corolla length of P. vulgaris also varied along an altitudinal gradient, and this variation strongly correlated with the local pollinator size but did not correlate with altitude itself. Furthermore, we found that the size match between the floral corolla length and bee proboscis length affected female and male plant fitness and the optimal size match (associated with peak fitness) was similar for the female and male fitness. Collectively, these results suggest that pollinator‐mediated selection influences spatial variation in the size of P. vulgaris flowers at a fine spatial scale.     

Do introduced honeybees (Apis mellifera,Hymenoptera) provide full pollination service to bird‐adapted Australian plants with small flowers? An experimental study of Brachyloma ericoides (Epacridaceae)

Abstract In many previous studies of the effects of introduced honeybees on Australian ecosystems, it has been assumed that floral morphology is a primary factor determining whether introduced honeybees will be effective in pollinating endemic plants. Although both honeybees and birds contacted stigmas and anthers of the small‐flowered Brachyloma ericoides (Epacridaceae), the exclusion of birds but not honeybees resulted in a significantly lower proportion of flowers producing capsules (12.3 ± 2 vs 21.0 ± 2%). This suggests that native birds contributed significantly to fruit set even though honeybees were much more frequent visitors to flowers (5–6 vs 0.7–2.5 times per day) and moved more frequently between plants (25 vs 12.2% of movements). Fruit set following exposure to birds and honeybees was very low compared with shrub species in general and may have been limited by the pre‐emptive removal of pollen by the 10% of honeybees that actively collected pollen.     

Beetle visitations,and associations with quantitative variation of attractants in floral odors of <Emphasis Type="Italic">Homalomena propinqua</Emphasis> (Araceae)

This study investigated floral visitations of two beetles, Parastasia bimaculata (Scarabaeidae) and Chaloenus schawalleri (Chrysomelidae), and examined associations between beetle visitations and variation in attractant traits, such as quantitative variations of attractants in floral odors and heat generation of spadices in Homalomena propinqua (Araceae). Observations showed P. bimaculata visited pistillate-phase inflorescences most frequently during heat generation, whereas C. schawalleri visited regardless of floral stages and heat generation. Chemical analyses of five dominant components of floral odors showed quantities of 2-butanol, veratrole, and α-pinene during the pistillate phase were the most abundant during all floral stages, and increased during heat generation. When testing combinations of these five authentic chemicals, some mixtures including 2-butanol or veratrole or both attracted both beetles, and veratrole attracted C. schawalleri. These results strongly suggested that the increased emission of floral odor attractants which accompanied heat generation influences floral visitations by P. bimaculata, but not by C. schawalleri. We therefore hypothesize that P. bimaculataaculata is a reliable pollinator, and that variation in attractant traits is a honest signal for P. bimaculata to seek rewards. In contrast, C. schawalleri can detect the signal even at low levels, and so visits inflorescences steadily during all floral stages.     

不同林龄杉木人工林土壤C:N:P化学计量特征及其与土壤理化性质的关系

为了阐明林龄对杉木人工林土壤理化性质及碳氮磷(C∶N∶P)生态化学计量特征的影响,在福建农林大学三明莘口林场选取4个林龄(4, 20, 24, 33a)的杉木人工林为研究对象,测定0—20,20—40,40—60 cm深度土壤的水分-物理性质、pH、总碳(TC)、全氮(TN)、全磷(TP)、全钾(TK),探讨它们随林龄的变化及其与C∶N∶P化学计量比之间的关系,为杉木人工林持续经营管理提供科学依据。结果表明:(1)随着林龄的增加,质量含水量、田间持水量和毛管孔隙度先减小后增加,在20 a达到最小,TN和TP也有相似的变化趋势,但在24 a林分的深层土壤达到最小,TC和TK保持不变;(2)随着林龄的增加,C∶N保持不变,C∶P和N∶P在24 a成熟林达到最大,但只在20—60 cm达到显著差异;(3)TC与多数水分-物理性质及3个生态化学计量比显著相关,质量含水量和孔隙度与C∶N∶P生态化学计量比均显著相关,TP与C∶P和N∶P显著负相关。土壤物理性质与土壤养分循环存在一定关联,有机质与土壤结构及养分平衡的调节有关,研究区杉木林发育过程中土壤腐殖质化进程较缓慢,24 a成熟林杉木的生长受到土壤磷的限制,在杉木速生阶段适当增施磷肥,保证林木的良好生长,促进土壤与植物的良性养分循环。