MORPHOLOGY AND ANATOMY OF FLORAL AND EXTRAFLORAL NECTARIES IN CAMPSIS (BIGNONIACEAE)

The genus Campsis (Bignoniaceae), with one New World and one Old World species, is unusual among temperate plants in having five distinct nectary sites. Multiple nectaries occur at all four of the extrafloral sites (petiole, calyx, corolla, fruit), representing an advanced strategy for ant attraction. The morphology and anatomy of the extrafloral nectaries in both species are uniform for the petioles, calyces, and young fruits; those on the outer corolla lobes are of slightly different forms. The generalized structure consists of one layer of basal cells, and a one- to two-layered secretory cup. Because of their small size, there is no vascular tissue in them. The large, vascularized (phloem only) floral nectary is an annular structure subtending the ovary.     

EFFECTS OF POLLINATION INTENSITY IN CAMPSIS RADICANS

I examined effects of pollination intensity on fruit, seed and seedling characteristics in Campsis radicans, and joint effects of pollen donor and pollination intensity on fruit production. Large pollen loads were more likely to initiate fruit production than small pollen loads, and the former fruits contained more seeds and a greater total seed mass. No further increases in seed number or mass occurred for pollen loads above 4,000 grains. The weight of individual seeds was unaffected by pollen load. Effects of pollen donor were generally larger than effects of pollen load, and fruit production from small loads of pollen from one donor were sometimes equal to fruit production from larger pollen loads from another donor. The ratio of pollen grains deposited to resultant seeds increased with pollen load, and several explanations are proposed. Seeds from heavy pollinations emerged better than seeds from light pollinations, but did not differ in speed of germination or in the performance of seedlings up to 126 days. The emergence differences are probably due to differing intensities of pollen tube competition.     

POLLINATION BIOLOGY OF LISTERA CORDATA (ORCHIDACEAE)

Pollination biology of northern California populations of Listera cordata (L.) R. Br. is presented. The flowers are allogamous, self-compatible, and protandrous. At anthesis, access to the stigma is blocked by the flap-like rostellum. When an insect contacts the touch-sensitive trigger hairs of the rostellum, the pollinia are suddenly released and become cemented to the insect's body. One to two days after the pollinia are removed, the rostellum rises and exposes the receptive stigma. Visitors are attracted to the flowers by their foul odor and minute quantities of nectar presented on the surface of the labellum. Pollination efficiency is high with 61–78% of flowers from several localities setting fruit. The most important and abundant pollinators of L. cordata at our study sites are fungus gnats (Sciaridae and Mycetophylidae). Other less frequently encountered pollinators are members of these families: Braconidae, Ichneumonidae, and Tipulidae. At present, there is no compelling evidence that the floral characteristics of L. cordata have evolved as a result of an evolutionary interaction with any specific group of insects. Nor is there any indication that the flowers are sapromyophilous, despite their putrid odor.     

POLLINATION AND THE FUNCTION OF FLORAL PARTS IN CHAMAECRISTA FASCICULATA (FABACEAE)

Chamaecrista fasciculata has a rigid upper petal that curves over nine of the ten anthers. The single anther is deflexed, as is the style. An experimental pollination study was designed to test the hypotheses that: 1) the rigid petal acts as flight guide to ensure pollination; and 2) the nine grouped anthers serve as fodder anthers, whereas the single deflexed anther functions in pollination. The rigid petal was removed from 97 flowers. Only 5% of the manipulated flowers set fruit in comparison to a fruit set of 47% for the control group. The results of the study support the flight guide hypothesis. Pollen from both sets of anthers is viable and germinates on receptive stigmas. A pollen flow experiment using powdered metals, backscatter scanning electron microscopy, and X-ray microanalysis revealed that there is no difference in the frequency of pollen distribution from either set of anthers. Therefore, this study refutes the fodder/pollinating anther hypothesis.     

POLLINATION BIOLOGY AND CHEMOTAXONOMY OF THE ECHINOCEREUS VIRIDIFLORUS COMPLEX (CACTACEAE)

The Echinocereus viridiflorus complex is morphologically distinct from other members of the genus. Temporal differences in flowering preclude natural interspecific hybridization between taxa of the E. viridiflorus complex and sympatric congeners. All taxa within the complex are xenogamous and self-incompatible. Considerable taxonomic disagreement exists among herbarium and field botanists with regard to species boundaries and infraspecific classification of the taxa constituting the E. viridiflorus complex. One author views the complex as consisting of a single species, E. viridiflorus while other authors additionally recognize E. chloranthus, E. russanthus, and E. davisii. Pollination in E. viridiflorus is by solitary halictid bees. Photographs taken with a filter transmitting long-wave ultraviolet (UV) light revealed two floral patterns among entities of the complex: entirely UV-absorbing tepals and tepals with UV-reflective outer segments and UV-absorbing centers (target-type blossoms). We suggest that UV-floral patterning increases pollinator visitation and efficiency. Six flavonol-O-glycosides and two dihydroflavonol-7-O-glycosides were localized in tepal tissue. No qualitative differences were seen either between differentially UV-patterned flowers or among E. viridiflorus, E. chloranthus, E. davisii, and E. russanthus. Flavonoid evidence indicates that these taxa are conspecific.     

REPRODUCTIVE BIOLOGY OF APIACEAE. I. FLORAL VISITORS TO THASPIUM AND ZIZIA AND THEIR IMPORTANCE IN POLLINATION

Based on the ready availability of nectar and pollen, and on the large numbers and great diversity of insect visitors, species in the Apiaceae have been labeled promiscuous. The distinction between floral visitors and effective pollinators, however, is extremely important and is rarely discerned. Floral visitation was documented for plants of nine populations in a comparative study of three species of the closely related apioid genera, Thaspium Nutt. and Zizia Koch. A pollinator importance index was calculated for each floral visitor using visitor abundance, pollen load composition and foraging behavior as its basic components. Results showed that, despite a high diversity of insect visitors, generally 1–4 species accounted for a minimum of about 74% of the pollinations in all populations. This specialization in pollination appears in part to be the result of an oligolectic relationship between Andrena ziziae (Hymenoptera; Andrenidae) and plants of the taxa studied, but solitary bees of Andrenidae, Colletidae and Halictidae in general were efficient and important pollinators. This study emphasizes that visitation records, when considered alone, effectively disguise specialization in the pollination system.     

THE POLLINATION BIOLOGY OF TUCKAHOE,PELTANDRA VIRGINICA (ARACEAE)

This study describes the specialized brood-site-based pollination system of Peltandra virginica Kunth (Araceae) with the chloropid fly Elachiptera formosa Loew and provides experimental evidence that gender-related changes in floral odor composition synchronize pollinator behavior with the blooming sequence. P. virginica is protogynous and does not self-pollinate because of a strong temporal separation in sexual function, and it is dependent upon insects for pollination because the spathe completely surrounds the spadix during the pistillate stage. Field observations conducted in central New Jersey and southeastern Pennsylvania showed that E. formosa is closely associated with P. virginica inflorescences. Within the floral chamber, E. formosa adults feed on pollen, mate, and find oviposition sites while the larvae complete their development. Although drosophilid and syrphid flies were collected and reared from inflorescences of both sexual stages, only E. formosa emerged from pistillate-stage inflorescences and adult E. formosa rapidly transferred fluorescent dye particles between inflorescences of both sexual stages in laboratory enclosures. These findings indicate that this fly species is the primary pollinator at our study locations. Field censuses demonstrated that although E. formosa visited P. virginica inflorescences of both sexual stages with equal frequency, the female flies preferentially oviposited within pistillate-stage inflorescences. Analysis of floral volatiles with gas chromatography-mass spectrometry showed that only the spathes emit the floral fragrance and that the composition of the floral volatiles changed during the transition from pistillate to staminate stage. A particularly noticeable change occurred in the emission ratio of the two primary floral odor components, 4,5,7-trimethyl-6,8-dioxabicyclo[3.2.1.]octane and an unidentified chemical analog with molecular weight of 142. The relationship between floral volatile composition and fly oviposition behavior was evaluated using sham inflorescences placed in the P. virginica study population. The sham inflorescences were constructed by covering spadices dissected from mature inflorescence buds with spathes excised from different-age inflorescences. After a 2-hr-long exposure period, sham inflorescences fitted with pistillate-stage spathes contained more E. formosa eggs than the sham inflorescences with staminate-stage spathes. However, the highest number of ovipositions occurred in sham inflorescences fitted with spathes excised from mature inflorescence buds. The fragrance emitted by these spathes was composed almost entirely of the two principal odor components. The decline in ovipositions observed in both real and sham inflorescences corresponded to an increase in the emission ratio of 4,5,7-trimethyl-6,8-dioxabicyclo[3.2.1.]octane to the molecular weight 142 analog. This suggests that gravid flies searching for oviposition sites used the ratio of the two primary floral volatile components as an inflorescence gender or age recognition cue.     

THE DELIMITATION OF BIGNONIACEAE AND SCROPHULARIACEAE BASED ON FLORAL ANATOMY,AND THE PLACEMENT OF PROBLEM GENERA

The delimitation of Bignoniaceae and Scrophulariaceae has long been a taxonomic problem. Several genera, including Paulownia, Schlegelia, Gibsoniothamnus, and Synapsis, have been variously placed in one or the other family. Differences between these two families have been noted with regard to the presence of endosperm, embryo and seed morphology, and placentation; however, the lack of comprehensive data on the distribution of such characters within these two families left the delimitation problem unsolved. A comprehensive study of floral anatomy confirmed a basic difference in the placentation of these two families, as well as a basic difference in gynoecial vascularization. Paulownia has a floral anatomy, embryo morphology, and seed morphology consistent with placement in Scrophulariaceae. While reminiscent of Bignoniaceae, Paulownia is not an intermediate genus linking the two families. Schlegelia and Gibsoniothamnus have a floral anatomy consistent with placement in Scrophulariaceae. Schlegelia also has a scrophulariaceous seed morphology. Considered anomalous in the Bignoniaceae, the Schlegelieae similarly are distinct in the Scrophulariaceae.     

小慈姑的开花状态、传粉机制与交配系统

在自然种群中对沼生植物小慈姑（Sagittaria potamogetifolia Merr.）的开花状态和传粉过程作了观测,并用同工酶遗传标记法定量估测了其3个种群中5个样本的异交率。该种为中心媒传粉为主的虫／风 媒兼性传粉机制,访花昆虫的飞行距离多在0-2m范围内,并与花序密度呈不显著的正相关。异交率（－／^／t）为50.0％－92.8％,表明该种为异交为主的混合交配系统（ixed mating system）。对雌雄同株的小慈姑而言,异交率与各种群花期植株密度无关,异交水平取决于种群中开花花序的两个或两个以上的植株所占比例。     

THE FLORAL BIOLOGY OF CASSIA DIDYMOBOTRYA AND C. AURICULATA (CAESALPINIACEAE)

The flowers of Cassia didymobotrya and C. auriculata have three types of fertile anthers that differ in orientation, size, shape, and pollen production. Flowers with right-handed or left-handed style deflections, i.e., enantiostylous flowers, occur in the same raceme. In Israel, both species are pollinated by pollen collecting buzzing females of Xylocopa pubescens. Vibrations of the carpenter bee are necessary for release of pollen from poricidal anthers and, in C. didymobotrya, also for absorption of pollen through the stigmatic opening into a cavity of the style tip. The largest anthers supply pollen for pollination. Pollen for consumption by the bee, on the other hand, appears to be supplied by all fertile anthers. This is in contrast to the accepted view of a complete separation of functions in the heteromorphic anthers in Cassia. The pollen/ovule ratio in C. didymobotrya was 55,200, and that in C. auriculata 32,000. Only 0.01% of the pollen produced by a flower was found in the stigmatic pollen load, and this fraction allows for high seed sets. From a comparison of the pollen production and utilization in the Cassia species and in certain distylous plants, it is concluded that heteranthery in Cassia does not confer an advantage in pollen economy. The high pollen/ovule ratios are explained by lack of nectar as well as by the large area on which pollen is deposited on the bee's body relative to the small size of the stigma. Enantiostyly in Cassia is considered as part of a pollination syndrome whose characteristics are outlined. The function of enantiostyly as an outbreeding strategy is discussed. In the two Cassia species, opportunities for self- and geitonogamous pollinations and self-compatibility minimize an effect of enantiostyly in promotion of outcrossing. Instead, it is proposed that style deflections may clear access of the pollinator to the anthers and protect female parts from injury by a vibrating heavy-bodied visitor.     

POLLINATION BIOLOGY,SEED PRODUCTION,AND POPULATION STRUCTURE IN QUEEN-OF-THE-PRAIRIE,FILIPENDULA RUBRA (ROSACEAE) AT BOTKIN FEN,MISSOURI

Queen-of-the-prairie, Filipendula rubra (Rosaceae), is a perennial, rhizomatous herb restricted to calcareous fens, spring seeps, and wet prairies, and is distributed in the northcentral United States from Iowa to New York. Because of the scarcity of adequate habitat, F. rubra is a rare and endangered species in several states. At Botkin Fen and other fens in Missouri, F. rubra possesses limited sexual reproduction. A study was conducted to determine causal mechanisms of its limited seed production that may be barriers to establishing this rare and endangered species in other fens in Missouri. A genetic-ecological hypothesis was supported. F. rubra was found to be self-incompatible. At Botkin Fen, most inflorescences produced very few seeds because most (85.4%) belonged to a single clone. Pollinators (Lassioglossum sp.) had a high probability of transporting pollen with the same self-incompatibility alleles. A few genetically different clones did produce significant quantities of seed. However, this seed, with its significant pool of genetic variability, was not recruited into the population. Apparently, competition from the dense sedges and grasses prevented the establishment of new seedlings.     

刺五加传粉生物学研究

本文报道刺五加Eleutherococcus seticosus(Rupr．et Maxim．)Maxim的花朵酬物、访花者类别、访花 者在花序上的访花行为、访花频率及传粉效果。主要结果如下:(1)每朵雄花可提供的报偿是58 000～ 81 000粒花粉,5.5～8.0 µl／d×2～4d花蜜。每朵雌花仅可提供4.0～8.0µl／d× 2～3 d花蜜,不能提 供花粉。每朵两性花可提供19 000～54 000粒花粉和7.0～10.0 µl／d× 2～6 d花蜜。(2)花粉提供者 (雄花和两性花)提供报偿的日期是开花后1～3天,花粉接受者(雌花和两性花在柱头外翻变白时)是 开花后5～7或7～9天,这进一步证实了剌五加雄蕊先熟。另外,花粉提供者提供花蜜的时间在一天中 是8:30至15:30,其高峰是9:00～15:00;花粉接受者提供花蜜的时间在一天中是10:30至16:30,其高 峰是11:00～16:00。这种时间差异可能是刺五加影响传粉者流向(从花粉提供者到花粉接受者)的关键 因素。(3)在刺五加花朵上记录到的访花昆虫有50余种,分别隶属于膜翅目、鳞翅目、鞘翅目、双翅目和 半翅目。在不同天气、不同生境、不同性别的植株记录到的访花频率及高峰时间不同:花粉提供者的 访花高峰早于花粉接受者的;雄株上的访花频率高于雌株的;两性株接受花粉时的访花频率高于提供花 粉时的;而天气越晴朗,刺五加地块(patch)越大,则访花频率越高;每日接受光照越早,访花高峰越早。 (4)从传粉效果看,刺五加种子的形成完全依赖于传粉昆虫的活动。其中,蜂类,如熊蜂、花蜂、切叶蜂、胡蜂和蜜蜂等是最有效的传粉者,其它昆虫如粉蝶、食蚜蝇、寄蝇和鳃角金龟等对传粉也有帮助。     

FLOWER COLOR POLYMORPHISM AND POLLINATION BIOLOGY OF PLATYSTEMON CALIFORNICUS BENTH. (PAPAVERACEAE)

Five genetically controlled flower color morphs in Platystemon californicus tend to occur in distinct geographic regions, suggesting regional selection of morphs. I examined the reproductive biology of P. californicus to determine whether color morphs are subject to selection due to differences in pollinator attractiveness. Plants of P. californicus have a high pollen: ovule ratio and are highly self-incompatible. Solitary bees (Andrenidae and Halictidae) are the most effective pollen vectors, but show inconsistent discrimination among color morphs. Platystemon californicus is unusual among annual self-incompatible plants in that wind is an important pollen vector. As a result of wind pollination, most polymorphic populations show no intermorph seed-set differences. Therefore, the geographic distribution of flower color morphs cannot be explained by differential attractiveness of color morphs to pollinators in different regions. Any selective value of color morphs must be due to linkage of flower color with as yet undetected morphologically or physiologically adaptive characters.     

FLORAL BIOLOGY OF NEPENTHES GRACILIS (NEPENTHACEAE) IN SUMATRA

Nepenthes gracilis, a dioecious carnivorous plant, has inconspicuous flowers lacking petals. Nectaries distributed on the upper surface of four sepals secrete dilute nectar (3%–12% sugar concentration) at night, but the nectar immediately disappears during the day by evaporation in the sunny environment of Sumatra. Male flowers have a higher nectar production rate but lower sugar concentration of nectar than female flowers. Flowers of both sexes were visited by pyralid moths at night and by calliphorid flies in the evening. Pollen was found attached on these insects visiting Nepenthes flowers. The pattern of nectar production of sepals is regarded as attracting nocturnal flying insects and avoiding ants, while the pitchers attract ants by nectar secreted on the pitcher rim.     

杭州石荠苎和石香薷（唇形科）的传粉生物学比较研究

杭州石荠苎（ＭｏｓｌａｈａｎｇｃｈｏｕｅｎｓｉｓＭａｔｓｕｄａｓ）和石香薷（Ｍ．ｃｈｉｎｅｎｓｉｓＭａｘｉｍ．）是一对种对,前零星分布于浙江,后广布于长江流域以南地区。杭州石荠苎以异花传粉为主,花粉／胚珠（Ｐ／Ｏ）为７５６＝－６１６３;石香薷以自花传粉为主,Ｐ／Ｏ＝１１０。     

POLLINATION BIOLOGY OF PHYSALIS VISCOSA VAR. CINERASCENS (SOLANACEAE)

Physalis viscosa var. cinerascens is a self-incompatible, herbaceous perennial which occurs in disturbed or open habitats in the south-central United States and eastern Mexico. The plants are low-growing and the pendant blossoms are often obscured by the foliage. In Oklahoma the species blooms from April through October. The flowers are yellow with dark purple or brown maculations in the center and a mat of white hairs in the throat. Nectar is produced in small quantities at the base of the ovary and pollen is present in copious amounts. The pollinators are solitary bees which visit the flowers for nectar and pollen. Perdita halictoides is the primary pollinator and is oligolectic on Physalis.