Preferences of pollinators and herbivores in gynodioecious Geranium sylvaticum

BACKGROUND AND AIMS: For the maintenance of gynodioecy (i.e. the coexistence of female and hermaphroditic plants), females need to compensate for the lack of pollen production through higher seed production or better progeny quality compared to hermaphrodites. In Geranium sylvaticum, females produce more seeds per flower than hermaphrodites. This difference in seed production might be modified by biological interactions with pollinators and herbivores that may favour one sex and thus affect the maintenance of gynodioecy. METHODS: Sexual dimorphism in flower size and flowering phenology, and in attractiveness to pollinators, pre-dispersal seed predators and floral herbivores were examined in natural populations of G. sylvaticum. KEY RESULTS: Pollinators preferred hermaphrodites 25 % more often than females in two of the three study populations, and floral herbivores attacked hermaphrodites 15 % more often than females in two of the six study populations. These preferences might be explained by the larger flower size of hermaphrodites. In contrast, seed predators did not prefer either sex. CONCLUSIONS: The data suggest that pollinator preference does not benefit females, whereas the higher floral herbivory of hermaphrodites might enhance the maintenance of females in G. sylvaticum. Thus, although the data support the view that ecological factors may contribute to the maintenance of gynodioecy, they also suggest that these contributions may vary across populations and that they may function in opposite directions.     

Floral nectaries, nectar production dynamics and chemical composition in six ipomoea species (convolvulaceae) in relation to pollinators

BACKGROUND AND AIMS: Floral nectaries and nectar features were compared between six Argentinian Ipomoea species with differences in their pollinator guilds: I. alba, I. rubriflora, I. cairica, I. hieronymi var. hieronymi, I. indica, and I. purpurea. METHODS: Pollinators were recorded in natural populations. The morpho-anatomical study was carried out through scanning electron and light microscopy. Nectar sugars were identified via gas chromatography. Nectar production and the effect of its removal on total nectar sugar amount were determined by using sets of bagged flowers. KEY RESULTS: Hymenopterans were visitors of most species, while hummingbirds visited I. rubriflora and sphingids I. alba. All the species had a vascularized discoidal nectary surrounding the ovary base with numerous open stomata with a species-specific distribution. All nectar samples contained amino acids and sugars. Most species had sucrose-dominant nectars. Flowers lasted a few hours. Mean nectar sugar concentration throughout the lifetime of the flower ranged from 34.28 to 39.42 %, except for I. cairica (49.25 %) and I. rubriflora (25.18 %). Ipomoea alba had the highest nectar volume secreted per flower (50.12 microL), while in the other taxa it ranged from 2.42 to 12.00 microL. Nectar secretion began as soon as the flowers opened and lasted for a few hours (in I. purpurea, I. rubriflora) or it was continuous during the lifetime of the flower (in the remaining species). There was an increase of total sugar production after removals in I. cairica, I. indica and I. purpurea, whereas in I. alba and I. rubriflora removals had no effect, and in I. hieronymi there was a decrease in total sugar production. CONCLUSIONS: The chemical composition, production dynamics and removal effects of nectar could not be related to the pollinator guild of these species. Flower length was correlated with nectary size and total volume of nectar secreted, suggesting that structural constraints may play a major role in the determination of nectar traits of these species.     

Floral biology and reproductive isolation by floral scent in three sympatric aroid species in French Guiana

We studied the reproductive biology of three sympatric Araceae species, Anthurium sagittatum, A. thrinax and Spathiphyllum humboldtii in French Guiana. The plants flowered simultaneously and were visited by scent‐collecting male euglossine bees, which were apparently their major pollinators. In total, each species was visited by 3–7 euglossine species, and 2–3 euglossine species accounted for at least 80% of all flower visits, with visits being plant species‐specific. Floral scent consisted of 6–10 main compounds, which made up 76–94% of the total amount of volatiles and were specific in these high amounts to each plant species. We suggest that the different floral scents lead to clear separation of the main pollinating euglossine species, providing a directed and efficient intraspecific pollen flow that results in high reproductive success. Since the simple floral (inflorescence) morphology of the studied plants does not support any morphological mechanisms to exclude visitors, as for example in euglossine‐pollinated perfume orchids, floral scent might be of major importance for the reproductive isolation and sympatric occurrence of these plants.     

Floral biology of Philippine morphospecies of the grape relative Leea (Leeaceae)

I observed the floral biology of three Leea morphospecies in a Philippine natural forest habitat. The red-flowered morphospecies Leea guineensis limits selfing through synchronized dichogamy, with male and female flowers temporally separated in the same inflorescence, whereas the two morphospecies of the white-flowered Leea indica might be prone to geitonogamous selfing. Light and soil pH are correlated with phenology. In addition to bees and flies, Leea is visited by wasps, butterflies, beetles, bugs and spiders.     

Chaotic Floral Phyllotaxis and Reduced Perianth in Achlys (Berberidaceae)

Among the 16 genera of the Berberidaceae Achlys is the only one with a reduced perianth, an irregular floral phyllotaxis, and variable stamen number. Early floral stages show an unstable (chaotic) arrangement of the organ primordia. Only the single carpel of the gynoecium has a more fixed position in that the placenta is formed in the adaxial half of the flower. The irregularities in the androecium may be caused by the lack of influence of a perianth on floral symmetry. On the other hand, the regular orientation of the carpel is perhaps due to the early polarity of the flower, whereby the abaxial half of the flower is larger (with further developed stamen primordia) at the time when carpel polarity is established.     

Pollination biology of Lilium japonicum var. abeanum and var. japonicum: evidence of adaptation to the different availability of diurnal and nocturnal pollinators

Floral traits such as color and size are highly diversified in lilies, but their adaptive significance remains uncertain. In the present study, we compared pollination processes between Lilium japonicum var. abeanum and var. japonicum to clarify how the two varieties are adapted to different pollinators. Lilium japonicum var. japonicum is known to be pollinated by moths, and we hypothesized from its flower traits that var. abeanum is pollinated by diurnal insects. Using waterproof digital cameras set to a recording interval of 10–30 s, we recorded flower visitors for 7–9 flowers of var. japonicum and 4–6 flowers of var. abeanum over 24‐h periods. We also recorded the number of versatile (T‐shaped) and rigid (I‐shaped) anthers per flower. For var. abeanum, we observed flowers at 05.00 and 17.00 hours to determine flower opening time and measured the intensity of floral scent using a metal‐oxide semi‐conductor odor sensor. Both diurnal and nocturnal insects visited flowers of the two varieties, but visitation of diurnal insects was more frequent in var. abeanum. Anthers of var. abeanum are usually rigid, as in many bee‐pollinated flowers, whereas those of var. japonicum are mostly versatile. Although flowers of var. japonicum are known to open in the evening, 32% of the flowers of var. abeanum started to flower during the day. Lilium japonicum var. abeanum emitted scent at night, with a maximum intensity at 20.00 hours, just as in var. japonicum. These findings suggest that the floral traits of the two varieties are adapted to the different relative availabilities of nocturnal versus diurnal pollinators. The flower of var. abeanum is more adapted to diurnal pollinators, but retains adaptations to nocturnal pollinators, particularly with regard to its scent.     

台闽苣苔(苦苣苔科)花部器官的形态发生

在扫描电镜下对台闽苣苔 (T. oldhamii (Hemsl.) Solereder)进行了花部器官形态发生的观察,为探索该类群的个体发育、类群间的系统发育关系和进化趋势提供依据.研究发现该属植物萼片、花冠和雄蕊发生式样均为五数花类型,它们各自来源于花原基上分化出来的萼片原基、花冠原基和雄蕊原基;花冠与雄蕊的两侧对称性与花冠上唇生长稍快和退化雄蕊原基发育迟滞相关;萼片原基的发生和发育的顺序是不一致的:萼片原基发生的式样为近轴中原基-远轴2原基-2侧原基,发育式样则为近轴中萼片-2侧萼片-远轴2萼片,花蕾时为镊合状排列.花冠裂片原基的发生和发育式样是一致的,即远轴中裂原基(下唇中裂片)-远轴2侧裂原基(下唇2侧裂片)-近轴2裂原基(上唇2裂片).花蕾期卷迭式为覆瓦状排列,从外向内:下唇中裂片-下唇2侧裂片-上唇2裂片或下唇2侧裂片-上唇2裂片-下唇中裂片.雄蕊原基与花冠裂片原基互生,前方雄蕊原基在发生上稍迟于后方雄蕊原基,后者与退化雄蕊原基几乎同时发生,但较小,并与近轴心皮(或柱头上唇)对生.将该属与玄参科(Scrophulariaceae)的地黄属( Rehmannia )、苦苣苔科(Gesneriaceae)的异叶苣苔属( Whytockia)和尖舌苣苔属(Rhynchoglossum )的花部器官比较发现,这四个属在这方面呈现出多样性和交叉.过去一直按子房室数和胎座类型划分玄参科(子房2室、中轴胎座)和苦苣苔科(子房1室、侧膜胎座)这一做法受到了质疑.     

Floral biology and pollinators of Trichocereus pasacana (Cactaceae) in Parque Nacional Los Cardones, Argentina]

Many columnar cacti are bat pollinated. It has been suggested that this kind of pollination would be more important in tropical than in temperate regions where flowers are open only one night. Thrichocereus pasacana produces big and resistant white flowers. We analyzed flower characteristics, floral cycle, stigmatic receptivity, nectar production, pollen presence and floral visitors in a T. pasacana population at National Park Los Cardones (Salta, Argentina) in November 1997. Flower features were constant between individuals of the population. Flowers start opening at evening and anthesis time is from 18 to 40 hs. The estigma was receptive throughout the floral cycle. Anther dehiscence occurs with flower opening. Nectar production was highest between 18 to 24 hs. Although T. pasacana are open during the night, floral visitors are diurnal. The most frequent was Xylocopa sp. In the study area, nectarivorous bats were not detected. The morphological features of T. pasacana flowers were similar but bigger compared to other columnar cacti. Anthesis time was also longer while nectar production was lower. T. pasacana pollination at National Park Los Cardones is done by bees.     

Crane flies and microlepidoptera also function as pollinators in Epidendrum (Orchidaceae: Laeliinae): the reproductive biology of E. avicula

Crane flies and microlepidoptera have been recorded as pollinators in unrelated orchid groups, but these insects have never been recorded in Epidendroideae, the most species‐rich orchid subfamily, which includes one of the most diverse genera among Orchidaceae, Epidendrum. Based on data on phenology, floral morpho‐anatomy, pollinators, pollination mechanisms and breeding system, the reproductive biology of E. avicula was studied in south‐eastern Brazil. Epidendrum avicula possess osmophores that produce a citric fragrance at night. The flowers attract Tipulidae flies and several families of microlepidoptera that drink the nectar produced in a tube formed by the adnation of the labellum and column. As is common in Epidendrum, after removing the pollinarium, both crane flies and micro‐moths get trapped by the proboscis, which frightens the insects and inhibits any possible intent to immediately visit another flower. The behavior of the pollinators on flowers, plus the retention of the anther cap by the pollinarium, results in a reduction in the occurrence of geitonogamy. Because E. avicula is self‐incompatible, the consequence of pollinator behavior and the floral mechanisms tend to reduce the pollen loss. As far as we know, this is the first study to report the reproductive biology of a species of Epidendroideae pollinated by crane flies and microlepidoptera. Based on more recent concepts of plant–pollinator interactions, although E. avicula is pollinated by several species belonging to two distinct orders, suggesting an unspecialized pollination system is involved, nectar‐seeking microlepidoptera and Tipulidae flies can be recognized as a single functional group.     

Floral nectar production and nectary anatomy and ultrastructure of Echinacea purpurea (Asteraceae)

BACKGROUND AND AIMS: In spite of the impressive species diversity in the Asteraceae and their widespread appeal to many generalist pollinators, floral-nectary ultrastructure in the family has rarely been investigated. To redress this, a study using Echinacea purpurea, a plant of horticultural and nutraceutical value, was undertaken. Nectar secretion of disc florets was compared with floral nectary ultrastructure taking into account nectar's potential impact upon the reproductive success of this outcrossing species. METHODS: Micropipette collections of nectar in conjunction with refractometry were used to determine the volume and nectar-sugar quantities of disc florets throughout their phenology, from commencement of its production to cessation of secretion. Light, scanning-electron and transmission-electron microscopy were utilized to examine morphology, anatomy and ultrastructure of nectaries of the disc florets. KEY RESULTS: Florets were protandrous with nectar being secreted from anthesis until the third day of the pistillate phase. Nectar production per floret peaked on the first day of stigma receptivity, making the two innermost whorls of open florets most attractive to foraging visitors. Modified stomata were situated along the apical rim of the collar-like nectary, which surrounds the style base and sits on top of the inferior ovary. The floral nectary was supplied by phloem only, and both sieve elements and companion cells were found adjacent to the epidermis; the latter participated in the origin of some of the precursor cells that yielded these specialized cells of phloem. Companion cells possessed wall ingrowths (transfer cells). Lobed nuclei were a key feature of secretory parenchyma cells. CONCLUSIONS: The abundance of mitochondria suggests an eccrine mechanism of secretion, although dictyosomal vesicles may contribute to a granulocrine process. Phloem sap evidently is the main contributor of nectar carbohydrates. From the sieve elements and companion cells, an apoplastic route via intercellular spaces and cell walls, leading to the pores of modified stomata, is available. A symplastic pathway, via plasmodesmata connecting sieve elements to companion, parenchyma and epidermal cells, is also feasible. Uncollected nectar was reabsorbed, and the direct innervation of the nectary by sieve tubes potentially serves a second important route for nectar-sugar reclamation. Microchannels in the outer cuticle may facilitate both secretion and reabsorption.     

Floral trait variation in fourCyclamen (Primulaceae) species

The purpose of this study is to examine floral trait variation between fourCyclamen species that show variation in their ability to reproduce in the absence of pollinators and in levels of inbreeding. Pollen and ovule production, pollen/ovule ratio, pollen volume, petal length and width, diameter of the corolla mouth, pistil and stamen length, and stigma-anther separation varied significantly between the four study species. Flower, pollen and ovule production, pollen volume and corolla size were generally highest inC. hederifolium, the species with the lowest level of inbreeding (mean F_is = 0.329), intermediate in both species with relatively high levels of pollinator-mediated inbreeding,C. repandum (mean F_is = 0.658) andC. creticum (mean F_is = 0.748), and lowest in the highly inbredC. balearicum (mean F_is = 0.930). The two species with the most different inbreeding coefficients,C. hederifolium andC. balearicum, had lower pollen-ovule ratios and shorter longevities of stigma receptivity and pollen viability thanC. creticum andC. repandum. These patterns of variation in floral traits are discussed in relation to the relatedness, pollination ecology and levels of inbreeding of the four species.     

Both biotic and abiotic factors influence floral longevity in three species of Epidendrum (Orchidaceae)

Extended flower longevity has been widely defended to occur as a consequence of low pollination activity. To test this hypothesis in Neotropical rainforests, we selected three species of Epidendrum that flower during rainy conditions and commonly exhibit low levels of insect visits. Experimental studies carried out in 15 populations distributed along a wide spatial gradient detected the presence of pollinator limitation in all populations. Pollinia removal without pollen deposition increased longevity, and thus extended longevity could apparently be considered as an ecological advantage for these orchids. Nevertheless, flower longevity was severely shortened by abiotic factors like precipitation and humidity. Comparison with other tropical orchids showed that flower longevity was higher even in species that flower during periods of more pollinator activity than the ones studied here, which is contrary to the hypothesis that pollinators are the only driving force for long‐lived flowers in Epidendrum. Thus, although long floral longevities are apparently favorable for the reproductive success of these species, the high maintenance costs experienced during rainy conditions are also an important factor constraining the selection of long‐lived flowers in Epidendrum.     

Floral development of Iris decora Wall. (Iridaceae)

The outer tepal and stamen primordia arise as secondary primordia on the outer tepal-stamenprimordia, which are formed on the floral apex. The inner tepal primordia are formed directly on the floral apex. All the floral appendages are initiated in the second tunica layer and are homologous with regard to their origin and early development. A short perianth tube is formed as a result of intercalary growth in the common bases of the tepal primordia. The intercalary growth in the fused bases of the floral appendages elevates the peripheral zone. The floral apex thus appears as a shallow cup. Further intercalary growth results in the formation of an inferior ovary. The ovules are initiated as outgrowths on placental ridges from the lateral ovary wall, the trilocular appearance being the result of secondary cohesion of the parietal placentae.     

Floral biology and reproductive ecology ofClusia nemorosa (Clusiaceae) in northeastern Brazil

Between March 1993 and May 1995 the floral biology and the phenology ofClusia nemorosa G. Mey. (Clusiaceae) were studied in two dioecious populations in Pernambuco State, and some observations were made in gynodioecious populations, that were encountered only in Bahia State. The species flowers from June to January/February. The flowers are bowl shaped, and last only one day. The floral reward is resin which is produced by staminodia in all types of flowers. The sex ratio was 1:1 for the studied populations. The pollen viability was approximately 98%. Apomixis was not observed. The rate of fruit set, in hermaphrodite individuals, under natural conditions and the sex ratio of the gynodioecious populations studied suggest that the hermaphrodite individuals may behave as males.Clusia nemorosa is melittophilous and was visited by 16 species of bees. The main pollinators wereEuplusia mussitans, E. iopyhrra, Eulaema cingulata, E. nigrita, Euglossa cordata andEuglossa sp.     

Floral biology and histochemical analysis of Vanilla edwallii Hoehne (Orchidaceae: Vanilloideae): an orchid pollinated by Epicharis (Apidae: Centridini)

The genus Vanilla is the most diverse in Vanilloideae, with ca 90 species distributed among tropical regions. Despite their economic importance, studies on pollination of Vanilla are very scarce and data on pollinators of species endemic to Brazil are lacking. Based on fieldwork and laboratory investigations, the floral biology of V. edwallii was studied. The pollinators and pollination process were recorded at the Serra do Japi reserve, state of São Paulo, southeastern Brazil, and the presence of floral reward was also investigated. Vanilla edwallii blooms in summer. The lateral inflorescences produce up to four pale green flowers. The white labellum is united to the base of the column forming a mentum. In the studied population V. edwallii is pollinated by Epicharis (Hoplepicharis) affinis, where the males exhibit a territorial behavior, defending flowers from other possible flower visitors. The pollen is deposited on the scutellum of bees when they abandon the flower. The mentum region is dry, suggesting no nectar production. The only secretory structures are osmophores dispersed on the inner surface of the lip responsible for production of a sweet fragrance, which together with color and morphology of flowers is related to bee attraction. The labellum is rich in mucilaginous cells, while the mucilaginous substance is retained inside the cells. The histochemical analysis also detected the presence of phenolic compounds and starch concentrated mainly at the adaxial surface of the lip and around the vascular bundles.     

Floral biology of the dioecious species Laurus nobilis L. (Lauraceae)

The present study examines the cytological, physiological, chemical and ecological characteristics of pollen and nectar offered by male and female flowers of the dioecious plant Laurus nobilis. The various phases of floral phenology and the insect pollinators were observed. We used cytological methods to determine anther, pollen and nectary structure. Nectar sugar composition was evaluated by HPLC. Pollen viability in time was compared with cytoplasmic and intine water content. Pollen presentation was found to be reversible by opening and closing of anther valves, determined by hydration of the mechanical layer of the anther. Pollen, covered by pollenkitt, was presented for dispersal for 3 consecutive days and during this time the intine and cytoplasm lost water and pollen viability diminished. At germination exine ruptured together with the outermost layer of the intine. Nectaries of male flowers were observed on the anther filament and on staminodes of female flowers. The nectar consisted almost entirely of sucrose and was more concentrated in male flowers. Secreted through stomata, nectar was presented in a thin layer. In the study area, the main pollinators (about half the total number of all visits) were hymenopterans. Pollen is of the recalcitrant type due to its high water content (>30%) but its viability is long-lasting because the intine is thick and stores water, keeping the cytoplasm of the vegetative cell hydrated and viable, and because anther valves may close under adverse conditions, protecting the pollen. Insects are attracted by male and female flowers similarly, males offer nectar and pollen, whilst females only nectar.     

海韭菜的花器官发生

运用扫描电镜（SEM）观察了海韭菜（Triglochin maritimum）的花器官发生发育过程。结果表明：海韭菜花发育是典型的单子叶植物发生模式,即两轮花被片、两轮雄蕊和两轮心皮以三基数轮状交替发生,花器官是以向心向顶的方式发生的,未发现“花被片—雄蕊复合原基”。 发育后期雄蕊和与之对生的花被片之间的共同基部可能是相继向上居间生长的结果。花被片轮和雄蕊轮二者之间在发育位置、时间和速率上存在差异,内轮花被片原基和外轮雄蕊原基的不同发育时间和发育速度使得在成熟花中内轮花被片位于外轮雄蕊的内方。观察结果不支持水麦冬属植物的花是退化（或压缩）的花序侧分枝等假花的观点。     

Floral thermogenesis of three species of Hydnora (Hydnoraceae) in Africa

Background and Aims

Floral thermogenesis occurs in at least 12 families of ancient seed plants. Some species show very high rates of respiration through the alternative pathway, and some are thermoregulatory, with increasing respiration at decreasing ambient temperature. This study assesses the intensity and regulation of respiration in three species of African Hydnora that represent the Hydnoraceae, an unusual family of holoparasitic plants from arid environments.

Methods

Long-term respirometry (CO₂ production) and thermometry were carried out on intact flowers of H. africana, H. abyssinica and H. esculenta in the field, and short-term measurements were made on floral parts during the protogynous flowering sequence.

Key Results

For H. africana, there was no temperature elevation in either the osmophores or the gynoecial chamber in any phase, and mass-specific respiration rates of the flower parts were low (maximum 8·3 nmol CO₂ g⁻¹ s⁻¹ in osmophore tissue). Respiration tracked ambient and floral temperatures, eliminating the possibility of the inverse relationship expected in thermoregulatory flowers. Hydnora abyssinica flowers had higher respiration (maximum 27·5 nmol g⁻¹ s⁻¹ in the osmophores) and a slight elevation of osmophore temperature (maximum 2·8 °C) in the female stage. Respiration by gynoecial tissue was similar to that of osmophores in both species, but there was no measurable elevation of gynoecial chamber temperature. Gynoecial chamber temperature of H. esculenta could reach 3·8 °C above ambient, but there are no respiration data available. Antheral tissue respiration was maximal in the male phase (4·8 nmol g⁻¹ s⁻¹ in H. africana and 10·3 nmol g⁻¹ s⁻¹ in H. abyssinica), but it did not raise the antheral ring temperature, which showed that thermogenesis is not a by-product of pollen maturation or release.

Conclusions

The exceptionally low thermogenesis in Hydnora appears to be associated with scent production and possibly gynoecial development, but has little direct benefit to beetle pollinators.Key words: Pollination biology, Hydnora, thermogenesis, respiration rate, temperature, flowers, insects