Sexual systems play important roles in angiosperm evolution and exhibit substantial variations among different floras. Thus, studying their evolution in a whole flora is crucial for understanding the formation and maintenance of plant biodiversity and predicting its responses to environmental change. In this study, we determined the patterns of plant sexual systems and their associations with geographic elements and various life-history traits in dry-hot valley region of southwestern China, an extremely vulnerable ecosystem. Of the 3166 angiosperm species recorded in this area, 74.5% were hermaphroditic, 13.5% were monoecious and 12% were dioecious, showing a high incidence of diclinous species. Diclinous species were strongly associated with tropical elements, whereas hermaphroditic species were strongly associated with temperate and cosmopolitan elements. We also found that hermaphroditism was strongly associated with showy floral displays, specialist entomophily, dry fruits and herbaceous plants. Dioecy was strongly associated with inconspicuous, pale-colored flowers, generalist entomophily, fleshy fruits, and woody plants, whereas monoecy was strongly associated with inconspicuous, pale-colored flowers, anemophily, dry fruits, and herbaceous plants. In addition, hermaphroditic species with generalist entomophily tended to flower in the dry season, whereas diclinous species with specialist entomophily tended to flower in the rainy season. However, independent of sexual systems, plants that produce dry fruits tended to flower in the rainy season and set fruits in the dry season, but the opposite pattern was found for fleshy fruit-producing plants. Our results suggest that in the dry-hot valleys, plant sexual systems are associated with geographic elements as well as various life-history traits that are sensitive to environmental change.

Floral bracts (bracteoles, cataphylls) are leaf-like organs that subtend flowers or inflorescences but are of non-floral origin; they occur in a wide diversity of species, representing multiple independent origins, and exhibit great variation in form and function. Although much attention has been paid to bracts over the past 150 years, our understanding of their adaptive significance remains remarkably incomplete. This is because most studies of bract function and evolution focus on only one or a few selective factors. It is widely recognised that bracts experience selection mediated by pollinators, particularly for enhancing pollinator attraction through strong visual, olfactory, or echo-acoustic contrast with the background and through signalling the presence of pollinator rewards, either honestly (providing rewards for pollinators), or deceptively (attraction without reward or even trapping pollinators). However, studies in recent decades have demonstrated that bract evolution is also affected by agents other than pollinators. Bracts can protect flowers, fruits, or seeds from herbivores by displaying warning signals, camouflaging conspicuous reproductive organs, or by providing physical barriers or toxic chemicals. Reviews of published studies show that bracts can also promote seed dispersal and ameliorate the effects of abiotic stressors, such as low temperature, strong ultraviolet radiation, heavy rain, drought, and/or mechanical abrasion, on reproductive organs or for the plants' pollinators. In addition, green bracts and greening of colourful bracts after pollination promote photosynthetic activity, providing substantial carbon (photosynthates) for fruit or seed development, especially late in a plant's life cycle or season, when leaves have started to senesce. A further layer of complexity derives from the fact that the agents of selection driving the evolution of bracts vary between species and even between different developmental stages within a species, and selection by one agent can be reinforced or opposed by other agents. In summary, our survey of the literature reveals that bracts are multifunctional and subject to multiple agents of selection. To understand fully the functional and evolutionary significance of bracts, it is necessary to consider multiple selection agents throughout the life of the plant, using integrative approaches to data collection and analysis.

Glandular trichomes are traditionally regarded as a defence against insect herbivores in plants. A comprehensive examination of their occurrence across an entire flora is essential for understanding how plant-herbivore interactions influence species abundance, traits, and diversity. In this study, we gathered data on the presence of glandular trichomes in a diverse flora comprising 7751 angiosperm species from the Hengduan Mountains region in southwestern China, testing the effects of plant organ, life form, leaf habit, phylogenetic position, and geographical element on the occurrence of glandular trichomes. Our results showed that 1021 species (13.2%) in the studied flora possessed glandular trichomes. Reproductive organs (88.0%) displayed a significantly higher occurrence of glandular trichomes compared to vegetative organs (59.1%). Within flowers, the calyx (81.1%) was the most defended, followed by the corolla (17.4%), and the gynoecium (12.5%). The androecium (1.8%) was the flower component least associated with glandular trichomes. Herbaceous species (14.0%) were more likely to have glandular trichomes than woody species (11.6%). For woody plants, leaf habit (deciduous/evergreen) had no significant effect on the presence of glandular trichomes on leaf blades, but had a significant effect on the occurrence of glandular trichomes on reproductive organs, with deciduous woody species (11.0%) displaying a higher prevalence compared to evergreen woody species (8.3%). Interestingly, the proportion of species with glandular trichomes was notably lower in the tropical elements (6.8%) compared to both temperate and cosmopolitan elements (15.3% and 15.6%, respectively). Our findings suggest that in the Hengduan Mountains region, the evolution of glandular trichomes exhibits distinct patterns among various life forms, plant organs, leaf habits, and geographical elements, while showing limited influence from phylogenetic relatedness, highlighting the remarkable evolutionary adaptability of this defensive plant trait.

Plants have long been thought to be less dependent on pollinators for seed production at higher ele-vations due to adverse pollination environments.However,recent research has yet to consistently support the generality of this expectation.In this study,we asked whether pollinator dependence de-creases along an elevational gradient and how it varies with various reproductive traits.To answer these questions,we quantified pollinator-plant associations and various reproductive traits for 112 flowering plants spanning a large elevational gradient(990-4260 m a.s.l.)in the Qinghai-Tibet Plateau.We found that flowering plants in the Qinghai-Tibet Plateau region are highly dependent on pollinators for seed production(76.2％ of seed production was contributed by animal pollinators and 44.6％ of plants would produce no seed without pollinator visitation).Contrary to our expectation,there was no significant elevational gradient in pollinator dependence index.Although the pollinator dependence index was not significantly correlated with pollen limitation,flower size,floral longevity,or reward type,it was correlated with compatibility status and flowering time.These findings indicate that pollinator depen-dence does not decrease along an elevational gradient in the Qinghai-Tibet Plateau.Our study also highlights the severe vulnerability of flowering plant seed production to pollinator declines under global change in the Qinghai-Tibet Plateau region,particularly for early-flowering or self-incompatible plants growing at higher elevations(e.g.,subnival belt).

The length of time a flower remains open and functional - floral longevity - governs important reproductive processes influencing pollination and mating and varies considerably among angiosperm species. However, little is known about large-scale biogeographic patterns and the correlates of floral longevity. Using published data on floral longevity from 818 angiosperm species in 134 families and 472 locations worldwide, we present the first global quantification of the latitudinal pattern of floral longevity and the relationships between floral longevity and a range of biotic and abiotic factors. Floral longevity exhibited a significant phylogenetic signal, and was longer at higher latitudes in both northern and southern hemispheres, even after accounting for elevation. This latitudinal variation was associated with several biotic and abiotic variables. The mean temperature of the flowering season had the highest predictive power for floral longevity, followed by pollen number per flower. Surprisingly, compatibility status, flower size, pollination mode, and growth form had no significant effects on flower longevity. Our results suggest that physiological processes associated with floral maintenance play a key role in explaining latitudinal variation in floral longevity across global ecosystems, with potential implications for floral longevity under global climate change and species distributions.

Specialized bracts of Himalayan 'glasshouse' plants are well known for adapting to diverse stresses in alpine environments, thus ensuring normal sexual reproduction. However, little information is available on how such specialized plants cope with the elevational increase in stress. In this study, we determine the elevational pattern of ultraviolet (UV) protection provided by the translucent bracts of Rheum nobile, a giant 'glasshouse' plant species, endemic to the high Himalayas, and examine its effect on pollen germination and pollen tube growth. Both UV-A and UV-B radiation in the open air increased with elevation, but their intensity beneath bracts remained constant with elevation, suggesting that the bracts of R. nobile growing at higher elevations have an increased ability to screen UV radiation. Enhanced UV-B radiation, equivalent to that experienced at higher elevations (4800 m a.s.l.) compared to that at lower elevations (4200 m a.s.l.), significantly reduced pollen germination and pollen tube growth. Pollen grains from plants at higher elevations were not more tolerant to UV-B radiation. UV absorbance and the content of flavonoids in bracts increased with elevation, but a similar trend was not found in either UV reflectance or adaxial trichome density, suggesting that bract UV absorbance may play a more active role than bract UV reflectance in helping plants cope with the increasing UV radiation. Our results indicate that the bracts of R. nobile have the ability to cope with enhanced UV radiation with increasing elevation through increased UV absorbance, thus protecting pollen grains from injury caused by higher levels of UV radiation, and consequently ensuring normal sexual reproduction in stressful high-alpine conditions.

Clonal reproduction (i.e., production of potentially independent offspring by vegetative growth) is thought to provide plants with reproductive assurance. Thus, studying the evolution of clonal repro-duction in local floras is crucial for our understanding of the adaptive mechanisms plants deploy in stressful environments such as alpine regions. In this study, we characterized clonal plant species in the subnival belt of the Hengduan Mountains (a global biodiversity hotspot with extreme environmental conditions in southwest China), in order to determine the effects of sex system, growth form, and ele-vational distribution on clonality. We compiled clonality data of angiosperm species belonging to 41 families in the subnival belt of the Hengduan Mountains using published information. Of the 793 species recorded in the region, 47.92％ (380 species) are clonal species. Both sex system and growth form had significant effects on the occurrence of clonal reproduction:unisexual species (79.79％) were more likely to be clonal than bisexual species (43.63％), and herbaceous species (51.04％) were more likely to be clonal than woody species (16.67％). Compared with non-alpine-endemic species (44.60％), alpine-endemic species (58.33％) showed a significantly higher proportion of clonal reproduction. Further logistic regression analysis showed a positive association between incidence of clonality and elevational range, indicating that species distributed at high elevations are more likely to be clonal. Furthermore, the elevational gradients in clonality were contingent on sex system or growth form. This study reveals that plants in the subnival belt of the Hengduan Mountains might optimize their probability of reproduction through clonal reproduction, a finding that adds to our growing understanding of plant's adaptations to harsh alpine environments.

Caryota obtusa is an endangered and economically important species of the Arecaceae. The complete chloroplast genome sequence of this species is a circular molecule of 159,882 bp in size, including a pair of inverted repeats with length of 27,271 bp, separated by a large single-copy (87,645 bp) region and a small single-copy region (17,695 bp). In total, there are 131 genes, encoding 79 protein-coding genes, 40 tRNAs, and 10 rRNA genes, in which 123 genes, 69 CDSs, 37 tRNAs, and 10 rRNAs are unique, respectively. Phylogenetic inference confirmed the monophyly of the Caryota genus and its delimitation in subfamily Coryphoideae.

Isotrema cangshanense X.X.Zhu, H.L.Zheng & J.S.Ma, a new species from western Yunnan, China, is described and illustrated here. It is similar to I. utriforme, I. forrestianum, I. cucurbitoides and I. obliquum The major differences between them are outlined and discussed. A detailed description, along with line drawings, photographs, habitat and distribution, as well as a comparison to morphologically similar species, is also provided. Meanwhile, the new taxon is assessed as Vulnerable (VU D2), according to the IUCN Red List criteria.

Corollas (or perianths), considered to contribute to pollinator attraction during anthesis, persist after anthesis in many plants. However, their post-floral function has been little investigated within a cost-benefit framework. We explored the adaptive significance of corolla retention after anthesis for reproduction in Fritillaria delavayi, a perennial herb endemic to the alpine areas of the Hengduan Mountains, southwestern China. We examined whether the persistent corollas enhance reproductive success during seed development. Persistent corollas increased fruit temperature on sunny days, and greatly decreased the intensity of ultraviolet-B/C (UV-B/C) radiation reaching fruits. When corollas were removed immediately after pollination, fecundity and progeny quality were adversely affected. Measurements of flower mass and size showed no further corolla growth during fruiting, and respiration and transpiration tests demonstrated that both respiration rate and transpiration rate of corollas were much lower during fruiting than during flowering, indicating a slight additional resource investment in corolla retention after anthesis. Thus, seed production by F. delavayi may be facilitated by corolla retention during seed development at only a small physiological cost. We conclude that corolla retention may be an adaptive strategy that enhances female reproductive success by having a protective role for ripening seeds in the harsh conditions at high elevation.

Aim We tested the elevational herbivory defence hypothesis, which predicts that plants from low elevations are better defended than plants from higher elevations. Location The Qinghai-Tibetan Plateau in south-western China-the world's highest plateau. Taxon Angiosperms. Methods We collated binary spinescence data for 10,622 angiosperm species ranging from 600 to 6,000 m a.s.l. Logistic regression was used to quantify the elevational pattern in spinescence. Because spinescence is thought to be especially effective against mammalian herbivores, we also quantified the association between spinescence and mammalian herbivore richness. Results We found a unimodal relationship between elevation and spinescence, with the highest proportion of spinescent species occurring at mid-elevations. This unimodal relationship was present in perennial herbs, shrubs and trees, but not in annual herbs. Herbivorous mammal richness also showed a unimodal elevational pattern. A positive association between herbivorous mammal richness and the incidence of spinescent species suggests that elevational variation in herbivore pressure from mammals might drive elevational variation in spinescence. Main Conclusions Our findings further call into question the elevational herbivory defence hypothesis and shed new light on the potential causes of elevational gradients in plant diversity.

Although there has been much experimental work on leaf colour change associated with selection generated by abiotic environmental factors and antagonists, the role of leaf colour change in pollinator attraction has been largely ignored. We tested whether whitening of the apical leaves subtending the inflorescences of Saururus chinensis during flowering enhances pollinator attraction, and whether re-greening of the white leaves after flowering increases carbon assimilation and promotes seed development. White leaves were removed or covered, and the effects of these manipulations on pollinator visitation and subsequent reproductive success were assessed. The net photosynthetic rates of leaves of different colour were measured and their photosynthetic contributions to seed development were evaluated. Saururus chinensis is able to self-pollinate autonomously, but depends largely on flies for pollination. White leaves had different reflectance spectra from green leaves, and white leaves attracted significantly more pollinators and led to significantly higher fruit and seed set. Although leaf whitening resulted in a reduction in photosynthetic capacity, it translated into only a small decrease in seed mass. When leaves had turned back from white to green after flowering their photosynthetic capacity was similar to that of 'normal' green leaves and promoted seed development. The reversible leaf colour change in S. chinensis appears to be adaptive because it enhances pollination success during flowering, with a small photosynthetic cost, while re-greening of these leaves after flowering helps to meet the carbon requirements for seed development.

AbstractStudying the drivers of host specificity can contribute to our understanding of the origin and evolution of obligate pollination mutualisms. The preference–performance hypothesis predicts that host plant choice of female insects is related mainly to the performance of their offspring. Soil moisture is thought to be particularly important for the survival of larvae and pupae that inhabit soil. In the high Himalayas, Rheum nobile and R. alexandrae differ in their distribution in terms of soil moisture; that is, R. nobile typically occurs in scree with well‐drained soils, R. alexandrae in wetlands. The two plant species are pollinated by their respective mutualistic seed‐consuming flies, Bradysia sp1. and Bradysia sp2. We investigated whether soil moisture is important for regulating host specificity by comparing pupation and adult emergence of the two fly species using field and laboratory experiments. Laboratory experiments revealed soil moisture did have significant effects on larval and pupal performances in both fly species, but the two fly species had similar optimal soil moisture requirements for pupation and adult emergence. Moreover, a field reciprocal transfer experiment showed that there was no significant difference in adult emergence for both fly species between their native and non‐native habitats. Nevertheless, Bradysia sp1., associated with R. nobile, was more tolerant to drought stress, while Bradysia sp2., associated with R. alexandrae, was more tolerant to flooding stress. These results indicate that soil moisture is unlikely to play a determining role in regulating host specificity of the two fly species. However, their pupation and adult emergence in response to extremely wet or dry soils are habitat‐specific.

A prerequisite for the evolutionary stability of pollinating seed-consuming mutualisms is that each partner benefits from the association. However, few studies of such mutualism have considered the benefit gained by the pollinators. Here, we determined how the pollinating seed-predators ensure the provisioning of their offspring in the recently discovered mutualism between Rheum nobile and Bradysia flies. The correlation between flower fate and fly oviposition was examined. Floral traits and patterns of variation in fruit abortion and fly oviposition were investigated to determine whether female flies exhibit preferences for particular flowers when laying eggs. Indole-3-acetic acid (IAA) was quantified to determine whether female flies manipulate host physiology. Flowers that flies oviposited on had a significantly lower probability of fruit abortion compared with intact flowers. Females did not exhibit oviposition preference for any of the floral traits examined. There was no significant correlation between fruit abortion and fly oviposition in terms of either flower position or timing of flowering. IAA concentrations in oviposited flowers were significantly higher than in intact flowers. Our results suggest that oviposition by the mutualistic seed-consuming pollinator Bradysia sp., greatly reduces the probability of fruit abortion of its host, R. nobile; this may be attributed to the manipulation of host physiology through regulating IAA levels.

AimsIt is generally accepted that visual displays and floral scent play important roles in communication between flowering plants and their pollinators. However, the relative role of visual and olfactory cues in pollinator attraction is largely unknown. In this study, we determined the roles of both types of cue in attracting pollinators to Cornus capitata, a medium sized tree with each capitulum surrounded by four large, white, petaloid bracts.MethodsPollinator observations and pollination experiments were conducted in a natural population; the inflorescences' visual and olfactory signals were characterized by spectral and chemical analyses; the responses of pollinators to visual and olfactory cues were tested using dual-choice behavioural bioassays; the relative roles of visual and olfactory cues in pollinator attraction were tested by comparing the responses of pollinators to inflorescences subjected to three experimental treatments (intact, all bracts removed, and capitulum removed) within the natural population.Important FindingsFor fruit set, C. capitata is entirely dependent on pollinators, with a bee, Anthophora sp., being the main pollinator. Bracts present high colour distance and green contrast against the leaves. Twelve volatile compounds in the floral scent were detected, most of which have previously been reported to be attractive to a broad spectrum of bee species. Behavioural bioassays showed that both, visual cues alone and olfactory cues alone, are attractive to pollinating bees. However, visual cues alone attracted significantly more approaches than olfactory cues alone, while olfactory cues alone elicited a significantly higher landing percentage than visual cues alone. The finding suggests that, in the C. capitata-Anthophora sp. interaction, visual cues are mainly used for location from long distances, while olfactory cues mainly aid landing from short distances. Our results indicate that different modalities of floral cues should be considered together to understand fully the communication between flowering plant and pollinators.

Environmental conditions are known to affect the reproductive fitness of insects, as do host plants. Although the highly specialized bracts of Himalayan 'glasshouse' plants are thought to be an adaptive morphology that allows the plant to reproduce in harsh alpine environments, little information appears to exist concerning the benefit of these bracts for the pollinating insects. In this study, we examine whether the semi-translucent bracts of Rheum alexandrae, a giant herb endemic to the alpine zone of the Hengduan Mountains in south-western China, enhance the reproductive fitness of its pollinator. Rheum alexandrae depends mainly on mutualistic seed-consuming Bradysia flies for pollination. Bracts increased interior temperature on sunny days, acted as a buffer against fluctuating air humidity, greatly decreased the intensities of ultraviolet B/C radiation and provided shelter from strong winds. Bract removal significantly decreased adult oviposition and offspring performance of pollinators during flowering and fruiting, respectively. Our results indicate that the bracts of R. alexandrae enhance the reproductive fitness of its pollinating seed-consuming mutualist in alpine environments, which may be attributed to the positive effects of the bracts on the interior microenvironment. (C) 2015 The Linnean Society of London.

In this study, the processes of pollination ecology of Fritillaria delavayi were investigated to document its reproductive characteristics. Some individuals of F. delavayi could produce seeds under bagging without emasculation (11%), but the rate was significantly lower than that of the natural control (87%). It is suggesting that pollination of F. delavayi largely depends on pollen vectors. Bombus sushikini was the only effective pollinator of F. delavayi and the visitation frequency was 0.003 time xXflower(-1) x min(-1). Flowering of F. delavayi in whole population lasted for 35 d and single flower for 11 d. Pollen viability and stigma receptivity lasted for 9 d and were relatively long compared with other Fritillaria genus plants. Consequently, bumblebee pollination and long floral longevity seem to be important for reproductive assurance of F. delavayi in harsh alpine environments.

Knowledge of how germination and seedling establishment respond to soil water condition is crucial for plant conservation under global warming and land-use changes. We tested the flooding and drought tolerance of two plant species with different occurrences along a soil water gradient by assessing seed germination, seedling survival, seedling growth, and root characteristics. In the high Sino-Himalayas, Rheum alexandrae typically occurs in wetlands, R. nobile in scree or open slope with well-drained soils. Seeds and seedlings of the two species were subjected to different soil water conditions in controlled greenhouse experiments. Seed germination in both species was inhibited by high soil water content; however, seeds of R. alexandrae were more tolerant to flooding, especially to submergence. Seedling survival, biomass accumulation, root diameter, and root porosity of R. alexandrae increased significantly with increased soil water content, but submergence was lethal for seedlings. Seedling survival, biomass accumulation, and root length of R. nobile increased significantly in response to reduced soil water content. These results indicate that in the two species, seed germination and seedling establishment in response to different soil water condition are habitat-specific. Because both species are susceptible to moderate changes in soil water condition, their species-specific requirements with respect to this factor should be a consideration when planning their conservation.