Optimal pollinator attraction strategies in Trollius ranunculoides Hemsl. (Ranunculaceae) at different altitudes: increased floral display or promotion of nectar output?

For alpine plant species, patterns of resource allocation to functional floral traits for pollinator attraction can be highly significant in adaptation to low pollinator abundance and consequent pollen limitation. Increased pollination can be achieved either through a larger floral display or production of more pollen rewards. In this study, variation in resource allocation to different components for pollinator attraction was studied along an altitudinal gradient in Trollius ranunculoides, an obligate self‐incompatible out‐crosser of the Qinghai–Tibet Plateau. We compared resource allocation to conspicuous yellow sepals (which mainly provide visual attraction) and degenerate petals (which provide the major nectar reward) between populations at four altitudes. Furthermore, we investigated the contribution of sepals and petals to pollinator attraction and female reproductive success in an experiment with sepal or petal removal at sites at different altitudes. At the level of single flowers, resource allocation increased to sepals but decreased to petals with increasing altitude. Consistent with these results, sepals contributed much more to visitation rate and seed set than petals, as confirmed in the sepal or petal removal experiment. Sepals and petals contributed to female reproductive success by ensuring visitation rate rather than visitation duration. To alleviate increasing pollen limitation with increasing altitude, resource allocation patterns of T. ranunculoides altered to favour development of sepals rather than petals. This strategy may improve pollination and reproductive success through visual attraction (sepal) rather than nectar reward (petal) over a gradient of decreasing pollinator abundance.     

基于同质园栽培的21种淫羊藿属植物花部特征的分类学意义

以采自中国多地并同质园栽培于吉首大学种质资源圃的21种淫羊藿为研究材料,通过观测这些物种的花梗长、花直径、内萼片长宽、蜜距长、雌雄蕊长、花药长、花柱长、胚珠数、花粉大小及花粉量等16个花部特征数量性状和花序类型、有无蜜距、花瓣颜色、内萼片颜色、内萼片与花瓣相对长度及花粉颜色6个质量性状,并对其分类学意义进行探讨。结果表明,(1)21种淫羊藿内轮萼片花瓣状,4枚花瓣除无距淫羊藿外均特化成蜜距,但花部形态种间存在显著差异,尤其体现于花直径、内萼片长宽、蜜距长、雌蕊长、胚珠数、花粉量、花粉直径等特征。(2)主成分分析显示花直径、内萼片长、花药长、胚珠数和花粉直径等性状信息负荷量较大,在该属分类中起主要作用。(3)根据花部特征聚类分析,物种间区分良好,21物种可分为4支:第1支花小,花瓣短于内轮萼片,囊状距或兜状距;第2支花较大,花瓣远长于内轮萼片,长距状;第3支花小,花瓣没有特化形成蜜距;第4支花较小,花丝伸长明显,可达10 mm,花蜜距呈钻状。(4)依据所观察到的花部形态特征编写了淫羊藿属21种植物花部特征分种检索表。研究认为,基于同质园栽培下的淫羊藿属物种的花部特征具有分类学意义,可以为该属分类提供一定依据。     

草乌传粉过程中的广告效应与回报物质研究

虫媒传粉植物与其传粉者之间的相互作用被认为是被子植物花多样性的一个重要选择压力。这种相互作用体现在植物对传粉者的吸引以及传粉者行为对花粉的转运两个方面。本项研究通过去除不同的花部器官研究了草乌(Aconitum kusnezoffii)对其传粉者的吸引, 并结合传粉者的访问行为和草乌花的生物学特性探讨了传粉过程与交配系统的适应。红光熊蜂(Bombus ignites)是草乌的有效传粉者。去除花萼片显著降低了红光熊蜂的访问频率, 但去除特化成蜜腺叶的花瓣并没有显著改变红光熊蜂的访问频率, 这表明草乌吸引红光熊蜂的主要结构是由5枚萼片组成的花部外观形态, 而非花瓣。花蜜是草乌提供给红光熊蜂的回报物质, 糖浓度为39.23%, 组氨酸浓度为0.25 μg/μL。草乌花较大、单花花期长、雌雄异熟、花粉寿命长, 是一个自交亲和但需传粉者传粉完成繁殖过程的异交物种。草乌花序是无限花序, 当上部的花处于雄性阶段时, 下部的花正好处于雌性阶段。而红光熊蜂在草乌花序上的访问顺序通常自下而上, 带来异交花粉为下部的花进行异花授精, 同时又带走上部花的花粉, 这就很好地促进了草乌的异交。草乌雄蕊自外向内逐渐成熟, 是一种有效的限制传粉者单次访问浪费大量花粉的花粉装配策略, 能提高植物通过花粉散布获得的雄性适合度。     

Sex‐specific floral attraction traits in a sequentially hermaphroditic species

• ● Many angiosperms are hermaphroditic and produce bisexual flowers in which male (pollen export) and female (stigma receptivity) functions are separated temporally. This sequential hermaphroditism may be associated with variation in flower size, color, or pattern, all of which may influence pollinator attraction. In this study, we describe variation in these traits across discrete functional sex stages within and between 225 greenhouse‐grown individuals of Clarkia unguiculata (Onagraceae). In addition, to identify the effects of floral phenotype on pollinator attraction in this species, we examine the effects of these floral traits on pollen receipt in ~180 individuals in an experimental field array.
• ● Petal area, ultraviolet (UV)‐absorbing nectar guide area, and blue and green mean petal reflectance differ significantly across the functional sex stages of C. unguiculata. Male‐ and female‐phase flowers display significantly different pollinator attraction traits. Petal and UV nectar guide area increase as flowers progress from male phase to female phase, while blue reflectance and green reflectance peak during anther maturation.
• ● In field arrays of C. unguiculata, female‐phase flowers with large UV nectar guides receive more pollen than those with small nectar guides, and female‐phase flowers with high mean blue reflectance values are more likely to receive pollen than those with low blue reflectance. Female‐phase flowers with green mean reflectance values that differ most from background foliage also receive more pollen than those that are more similar to foliage. These findings indicate that components of flower color and pattern influence pollen receipt, independent of other plant attributes that may covary with floral traits. We discuss these results in the context of hypotheses that have been proposed to explain sex‐specific floral attraction traits, and we suggest future research that could improve our understanding of sexual dimorphism in sequentially hermaphroditic species and the evolution of features that promote outcrossing.

     

Floral ontogeny in Dipterygeae (Fabaceae) reveals new insights into one of the earliest branching tribes in papilionoid legumes

Flowers of Dipterygeae (Fabaceae, Papilionoideae) exhibit an unusual petaloid calyx. The two adaxial sepals are large and petaloid, and the three abaxial sepals form a three‐toothed lobe. The goal of this study was to elucidate the ontogenetic pathways of this peculiar calyx in light of the floral development of the three genera that comprise the tribe. Floral buds of Dipteryx alata, Pterodon pubescens and Taralea oppositifolia were analysed using scanning electron microscopy and light microscopy. The order of bracteole and sepal initiation varies among the species. The androecium is asymmetric. The carpel cleft is positioned to the right or to the left, and is opposite the adaxial antepetalous stamen. The peculiarity of the calyx becomes noticeable in the intermediate stages of floral development. It results from the differential growth of the sepal primordia, in which the abaxial and lateral primordia remain diminutive during floral development, compared with the adaxial ones that enlarge and elongate. Bracteoles, abaxial sepals, petals and anthers are appendiculate, except in T. oppositifolia, in which the appendices were not found in bracteoles or anthers. These appendices comprise secretory canals or cavities. Considering that the ontogenetic pathway for the formation of the petaloid calyx is similar and exclusive for Dipterygeae, it might be a potential synapomorphy for the group, with the presence of secretory canals in the appendices of abaxial and lateral sepals and petals. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 529–550.     

蓝堇草属(毛茛科)花形态发生的扫描电子显微镜观察

花的发生和发育过程研究可以发现早期进化的轨迹,为系统发育的研究提供重要线索。蓝堇草属(Leptopyrum)为毛茛科唐松草亚科一单种属,仅包含蓝堇草一种,其花的发生和发育过程仍为空白。为了深入理解唐松草亚科乃至毛茛科花发育多样性和演化规律,该文运用扫描电子显微镜(SEM)观察了蓝堇草各轮花器官的形态发生和发育过程。结果表明:该属植物所有的萼片、花瓣、雄蕊和雌蕊均为螺旋状发生,花器官排列式样也为螺旋状; 5枚萼片原基宽阔,5枚花瓣原基圆球形、位于萼片原基的间隔,且在后期表现为延迟发育现象,雄蕊原基较小、为圆球形; 花瓣原基和雄蕊原基连续发生,无明显的时空间隔,但与萼片原基有时空间隔; 心皮原基为马蹄形对折,柱头组织由单细胞乳突组成; 胚珠倒生、具单珠被。该属花器官螺旋状排列、胚珠具单珠被在唐松草亚科中是独有的性状,花发育形态学证据支持了该属的特殊性。     

长角凤仙花（凤仙花科）的花器官发生和发育

以不同发育时期的长角凤仙花Impatiens longicornuta Y.L.Chen（凤仙花科Balsaminaceae）为材料,利用扫描电镜技术观察了其花器官的分化及其发育过程。长角凤仙花为两侧对称花,具2枚侧生萼片,唇瓣囊状,旗瓣具鸡冠状突起,雄蕊5枚,子房上位,5心皮5室。其花器官分化顺序为向心式,萼片—花瓣—雄蕊—雌蕊原基。2枚侧生萼片先发生,然后近轴萼片（即唇瓣）原基和2枚前外侧萼片原基近同时发生;但是这3枚萼片原基的发育不同步,远轴的2枚前外侧萼片原基的发育渐渐滞后,然后停止发育,最后渐渐为周围组织所吸收,直至消失不见。花瓣原基中,旗瓣原基最先发生,4个侧生花瓣原基相继成对发生,且之后在基部成对愈合形成翼瓣;5枚雄蕊原基几乎同时发生,5个心皮原基轮状同时发生。本文结果支持凤仙花属植物为5基数的花,并进一步证实了唇瓣的萼片来源;此外,研究结果表明花器官早期发育资料对植物系统与进化研究具有重要参考价值。     

Floral ontogeny in legume genera Petalostylis, Labichea, and Dialium (Caesalpinioideae: Cassieae), a series in floral reduction

Floral ontogeny of taxa of two subtribes (Labicheinae, Dialiinae) of caesalpinioid tribe Cassieae, characterized by reduced number of floral organs, was compared. All three taxa studied are distichous; Petalostylis labicheoides flowers are solitary in leaf axils, Labichea lanceolata has few-flowered racemes, and Dialium guineense has numerous-flowered cymes. The first sepal primordium in each is initiated abaxially and nonmedianly. Order of organogenesis in Petalostylis is: five sepals bidirectionally, five petals and carpel simultaneously, then five stamens bidirectionally, starting abaxially. The order in Labichea is: five sepals helically (one lagging in time), five petals unidirectionally starting abaxially, the carpel and petals concurrently, then two stamens successively, starting laterally. Order in Dialium is: five sepals bidirectionally, the single petal adaxially, and lastly the carpel and two stamens concurrently. Specializations include (1) reduction of the five sepals to four by fusion in Petalostylis and Labichea; (2) reduction of petal number to one in Dialium; (3) reduction of stamen number to two in Labichea and Dialium, and reduction of functional stamens to three in Petalostylis; and (4) an elaborate, late-developing style in Petalostylis. Floral asymmetry, another specialization, characterizes Labichea, expressed by dissimilar stamens, while the other genera have zygomorphic flowers. Floral ontogenies are compared with other taxa of Cassieae.     

The development of pistillate and perfect florets in Xeranthemum squarrosum (Asteraceae)

The formation of capitulum inflorescence with two different types of floret is an interesting issue in floral biology and evolution. Here we studied the inflorescence, floral ontogeny and development of the everlasting herb, Xeranthemum squarrosum, using epi‐illumination microscopy. The small vegetative apex enlarged and produced involucral bracts with helical phyllotaxy, which subtended floret primordia in the innermost whorl. Initiation of floret primordia was followed by an acropetal sequence, except for pistillate peripheral florets. The origin of receptacular bracts was unusual, as they derived from the floral primordia rather than the receptacular surface. The order of whorl initiation in both disc and pistillate flowers included corolla, androecium and finally calyx, together with the gynoecium. The inception of sepals and stamens occurred in unidirectional order starting from the abaxial side, whereas petals incepted unidirectionally from the adaxial or abaxial side. Substantial differences were observed in flower structure and the development between pistillate and perfect florets. Pistillate florets presented a zygomorphic floral primordium, tetramerous corolla and androecium and two sepal lobes. In these florets, two sepal lobes and four stamen primordia stopped growing, and the ovary developed neither an ovule nor a typical stigma. The results suggest that peripheral pistillate florets in X. squarrosum, which has a bilabiate corolla, could be considered as an intermediate state between ancestral bilabiate florets and the derived ray florets.     

A flower with several secretions: anatomy,secretion composition,and functional aspects of the floral secretory structures of Chelonanthus viridiflorus (Helieae—Gentianaceae)

Floral secretory structures have been reported for Gentianaceae; however, morphoanatomical studies of these glands are rare. We described the development and secretory activity of the colleters and nectaries throughout the floral development of Chelonanthus viridiflorus. We collected flower buds, flowers at anthesis, and fruits to be investigated using light and scanning electron microscopy. We performed histochemical tests on the secretion of colleters and used glycophyte to confirm the presence of glucose in nectar. Colleters are located on the ventral surface of sepals and nectaries occur in four regions: (i) the dorsal and (ii) ventral surfaces of sepals; (iii) apex of petals; and (iv) base of ovary. The colleters have a short peduncle and a secretory portion with homogeneous cells. They are active in flower buds and secrete polysaccharides and proteins. In flowers at anthesis, they begin to senescence presenting protoplast retraction, cell collapse, and lignification; these characteristics are intensified in fruit. The nectaries of sepals and petals have two to five cells surrounding a central cell through which the secretion is released. Nectaries are numerous, forming a nectariferous area on the dorsal surface of sepals, like that observed on petals, and can form isolated units on the ventral surface of sepals. They are active from flower buds to fruits. A region with secretory activity was identified at the base of the ovary. The secretion of colleters acts in the protection of developing organs, while nectaries are related to defenses against herbivores and the supply of nectar to potential robbers or pollinators.

     

FLORAL DEVELOPMENT IN CAESALPINIA (LEGUMINOSAE)

Utilizing scanning electron microscopy, we studied the early floral ontogeny of three species of Caesalpinia (Leguminosae: Caesalpinioideae): C. cassioides, C. pulcherrima, and C. vesicaria. Interspecific differences among the three are minor at early and middle stages of floral development. Members of the calyx, corolla, first stamen whorl, and second stamen whorl appear in acropetal order, except that the carpel is present before appearance of the last three inner stamens. Sepals are formed in generally unidirectional succession, beginning with one on the abaxial side next to the subtending bracts, followed by the two lateral sepals and adaxial sepal, then lastly the other adaxial sepal. In one flower of C. vesicaria, sepals were helically initiated. In the calyx, the first-initiated sepal maintains a size advantage over the other four sepals and eventually becomes cucullate, enveloping the remaining parts of the flower. The cucullate abaxial sepal is found in the majority of species of the genus Caesalpinia. Petals, outer stamens, and inner stamens are formed unidirectionally in each whorl from the abaxial to the adaxial sides of the flower. Abaxial stamens are present before the last petals are visible as mounds on the adaxial side, so that the floral apex is engaged in initiation of different categories of floral organs at the same time.     

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

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

Janusia anisandra (A. Juss.) Griseb (Malpighiaceae): The first case of enantiostyly in oil-flower species

Enantiostyly is a floral polymorphism characterized by style deflection to the right and left, which results in flowers with mirror images. To date, enantiostyly occurs only in species that offer pollen or nectar as floral resource. Here we describe the first case of enantiostyly in an oil-flower species. Janusia anisandra (A. Juss.) Griseb (Malpighiaceae) shows right and left-styled flowers in individuals occurring in an area of tropical dry forest in Northeastern Brazil. We described the floral morphology, investigate the reproductive system, and made focal observations of flower visitors behaviour. J. anisandra has morphological traits that are part of the floral conservatism found in the neotropical species of Malpighiaceae, which preserves features that are decisive for the interaction with oil-collecting bees of the Centridini tribe, performing a highly specialized pollination system. The six stamens (two major and four smaller), located in the center of the flower, characterizes heteranthery, an intrafloral polymorphism, in which the stamens differ in shape, size, and function and is commonly associated with enantiostyly. We identified the two floral morphs in the same individual, characterizing a monomorphic enantiostyly; each of the morphs differs by lateral displacement of the style to the right or to the left, always accompanied by one larger stamen. The species is moderately self-compatible. Four species of solitary bees of the genus Centris were registered visiting the two morphs of J. anisandra flowers. Centris aenea and C. fuscata were the main pollinators, performing 65% of the visits. This new discovery of a classical monomorphic enantiostyly in J. anisandra opens the way for investigations in the evolution of this system in oil-secreting flowers.     

NORMAL FLORAL ONTOGENY AND COOL TEMPERATURE-INDUCED ABERRANT FLORAL DEVELOPMENT IN GLYCINE MAX (FABACEAE)

Floral onset in soybean (Glycine max cv. Ransom) is characterized by precocious initiation of axillary meristems in the axils of the most recently initiated leaf primordium. During floral transition, leaf morphology changes from trifoliolate leaf with stipules, to a three-lobed bract, to an unlobed bract. Soybean flowers initiated at 26/22 C day/night temperatures are normal, papilionaceous, and pentamerous. Sepal, petal, and stamen whorls are initiated unidirectionally from the abaxial to adaxial side of the floral apex. The median sepal is located abaxially and the median petal adaxially on the meristem. The organogeny of ‘Ransom’ flowers was found to be: sepals, petals, outer stamens plus carpel, inner stamens; or, sepals, petals, carpel, outer stamens, inner stamens. The outer stamen whorl and the carpel show possible overlap in time of initiation. Equalization of organ size occurs only within the stamen whorls. The sepals retain distinction in size, and the petals exhibit an inverse size to age relationship. The keel petals postgenitally fuse along part of their abaxial margins; their bases, however, remain free. Soybean flowers initiated at cool day/night temperatures of 18/14 C exhibited abnormalities and intermediate organs in all whorls. The gynoecium consisted of one to ten carpels (usually three or four), and carpel connation varied. Fusion of keel petals was often lacking, and stamen filaments fused erratically. Multiple carpellate flowers developed into multiple pods that were separate or variously connate. Intermediate type organs had characteristics only of organs in adjacent whorls. These aberrant flowers demonstrate that the floral meristem of soybean is not fixed or limited in its developmental capabilities and that it has the potential to produce alternate morphological patterns.     

Pollination biology in Roepera (Zygophyllaceae): How flower structure and shape influence foraging activity

The foraging behavior of bees is a complex phenomenon that depends on numerous physical features of flowers. Of particular importance are accessibility of floral rewards, floral proportions, symmetry and orientation. The flowers of Roepera are characterized by the presence of staminal scales (SS), which play an important role in nectar protection. We studied two species of Roepera with different symmetry and flower orientation, which are mainly visited by honeybees (Apis mellifera). We aimed to show how the foraging behavior of honey bees is affected by the function of SS, floral symmetry and orientation. The foraging behavior was documented by video photography. Handling time, access to nectar, percentage of pollen/nectar foraging, percentage of pollen contact and pollen deposition site on the honey bee's body were assessed. The morphometric features of the honey bees and flowers were analyzed. We found that the SS restricted pollinator access to nectar. Our results indicated consistency of visitation patterns in zygomorphic, laterally oriented flowers of R. fuscata versus random patterns in actinomorphic, diversely oriented flowers of R. leptopetala. The relative proportions of SS and proboscis length appear to be crucial for the success of pollinators. The directionality of the honey bees' movement, together with the different positioning of reproductive organs, plays an important role in the accuracy of pollen transfer and pollination efficiency.     

蝴蝶兰PhalPI基因的克隆及在花器官突变体中的表达分析

为深入研究兰科植物花器官发育的调控机理,从蝴蝶兰花瓣中克隆了一个B类MADS-box转录因子PhalPI（GenBank登录号为KY020416）。序列分析表明,该基因的cDNA全长为944 bp,含完整的开放阅读框,可编码210个氨基酸,属于BGLO/PI蛋白家族,与蝴蝶兰属的PhPI10和PeMADS6基因关系最近;表达模式分析表明,PhalPI基因在生殖器官中表达,在营养器官中不表达,在授粉后的子房中,该基因的表达水平降低。在5种花器官突变体中,PhalPI基因在萼片唇瓣化突变体的萼片和蕊柱中表达水平明显升高;在雄蕊花瓣化突变体的萼片和侧瓣中表达水平降低,在其唇瓣和蕊柱中显著升高;在侧瓣合柱化突变体的蕊柱中,PhalPI基因的表达也发生了显著升高;PhalPI基因表达的改变与花器官形态的突变相关;而在侧瓣唇瓣化和侧瓣花药化突变体中,PhalPI基因的表达水平没有变化。推测该基因在决定蝴蝶兰侧瓣和唇瓣的发育中起重要的调控作用。     

A SCANNING ELECTRON MICROSCOPIC STUDY ON THE INITIATION AND DEVELOPMENT OF FLORAL ORGANS OF BRASSICA NAPUS (CV. WESTAR)

The initiation and development of the floral organs of Brassica napus L. (cv. Westar) were examined using the scanning electron microscope. After transition of the vegetative apex into an inflorescence apex, flower primordia were initiated in a helical phyllotactic pattern. The sequence of initiation of the floral organs in a flower bud was that of sepals, stamens, petals and gynoecium. Of the four sepal primordia, the abaxial was initiated first, followed by the two lateral and finally the adaxial primordium. The four long stamens were initiated simultaneously in positions alternating with the sepals. The two short stamens were initiated basipetal to and outside the long stamens, and opposite the lateral sepals. The petals arose on either side of the two short stamens and the gynoecium was produced from the remainder of the apex. During development, the sepal primordia curved sharply at the tips and tightly enclosed the other organs. Stamen primordia developed tetralobed anthers at an early stage while filament elongation occurred just prior to anthesis. A unique pattern of bulbous cells was present on the abaxial surface of the anther. Growth of petal primordia lagged relative to the other floral organs but expansion was rapid prior to anthesis. The gynoecium primordium was characterized by an invagination early in development. At maturity, there was differentiation of a papillate stigma, an elongated style and a long ovary marked externally by sutures and divided internally by a septum. Distinct patterns of cuticular thickenings were observed on the abaxial and adaxial surfaces of the petals and stamens and on the surface of the style. The patterns were less obvious on the sepals and ovary. Stomata were present on both surfaces of the mature sepals, on the style and restricted areas on the abaxial surface of the anthers and nectaries but were absent from the petals, the adaxial surface of the stamens and the ovary. No hairs were present on any of the floral organs.     

Notes on the floral morphology in Vivianiaceae (Geraniales)

The functional floral morphology of the three genera of Vivianiaceae (= Ledocarpaceae, Geraniales), Rhynchotheca, Viviania and Balbisia, is compared. Likely pollination mechanisms are inferred from morphology and field observations. The flowers of Viviania are nectariferous and apparently zoophilous with nectar as the (primary) pollinator reward. Balbisia has pollen flowers without nectaries, its showy corolla indicates that it is also zoophilous with pollen as sole pollinator reward; bees were observed as flower visitors. One taxon (B. gracilis) may be anemophilous. Rhynchotheca has flowers without petals, with large, pendulous anthers and lacks nectaries. It shows synchronous mass flowering in its natural populations and is evidently anemophilous. A comparison with other Geraniales shows that nectar flowers with small anthers are likely the ancestral condition in Vivianiaceae. This suggests that the pollen flowers with larger anthers of Balbisia and Rhynchotheca may represent an apomorphic condition. The documentation of pollen flowers and anemophily in Vivianiaceae expands the range of known floral and pollination syndromes in Geraniales.     

Pollination biology of the sclerophyllous shrub Pultenaea villosa Willd. (Fabaceae) in southeast Queensland, Australia

The pollination biology of the common shrub Pultenaea villosa Willd. was examined in a subtropical dry sclerophyll forest in eastern Australia. We determined floral phenology and morphology, the timing of stigma receptivity and anther dehiscence, nectar availability, the plant breeding system, and flower visitors. The shrub's flowers are typical zygomorphic pea flowers with hidden floral rewards and reproductive structures. These flowers require special manipulation for insect access. A range of insects visited the flowers, although bees are predicted to be the principle pollinators based on their frequency on the flowers and their exclusive ability to operate the wing and keel petals to access the reproductive structures. Nectar and pollen are offered as rewards and were actively collected by bees. Nectar is offered to visitors in minute amounts at the base of the corolla. In Toohey Forest, P. villosa flowers in spring and is the most abundant floral resource in the understory of the forest at this time. The breeding system experiment revealed that P. villosa requires outcrossing for high levels of seed set and that the overlap of stigma receptivity and pollen dehiscence within the flower suggests the potential for self-incompatibility.     

Trends in evolution of floral ontogeny in Cassia sensu stricto,Senna, and Chamaecrista (Leguminosae: Caesalpinioideae: Cassieae: Cassiinae); a study in convergence

Distinctions in floral ontogeny among three segregate genera (Cassia sensu stricto, Chamaecrista, and Senna) of Cassia L. support their separation. In all species studied, the order of floral organ initiation is: sepals, petals, antesepalous stamens plus carpel, and lastly antepetalous stamens. Sepal initiation is helical in all three genera, which however differ in whether the first sepal is initiated in median abaxial position (Senna), or abaxial and off-median (Cassia javanica), a rare character state among legumes. Order of petal initiation varies: helical in Senna vs. unidirectional in Cassia and Chamaecrista. Both stamen whorls are uniformly unidirectional. Intergeneric ontogenetic differences occur in phyllotaxy, inflorescence architecture, bracteole formation, overlap of initiation among organ whorls (calyx/corolla in Cassia; two stamen whorls in Chamaecrista), eccentric initiation on one side of a flower, anther attachment, anther pore structure, and precocious carpel initiation in Senna. The asymmetric corolla and androecium in Chamaecrista arise by precocious organ initiation on one side (left or right). The poricidal anther character can result from differing developmental pathways: lateral slits vs. sealing of lateral sutures; clasping hairs vs. sutural ridges; terminal pores (one or two) vs. none; and clamp layer formation internally that prevents lateral dehiscence. Genera differ in corolla aestivation patterns and in stigma type. Convergence is shown among the three genera, based on intergeneric dissimilarities in early floral ontogeny (floral position in the inflorescence, bracteole presence, position of the first sepal initiated, order of petal initiation, asymmetric initiation, overlap between whorls, anther morphology, and time of carpel initiation) resulting in similarities at anthesis (showy, mostly yellow salverform flowers, heteromorphic stamens, poricidal anther dehiscence, bee pollination, and chambered stigma).