Comparative floral biology of Rhynchospora ciliata (Vahl) Kukenth and R. pubera (Vahl) Boeckeler (Cyperaceae): the role of white involucral bracts in attracting pollinating insects

Although Cyperaceae are considered anemophilous, some species exhibit features that are attractive to pollinators, such as the white UV‐reflecting involucral bracts of Rhynchospora ciliata. But how effective are these conspicuous adaptations? To address this question, we tested the hypothesis that species such as R. ciliata are visited by greater numbers of pollinating insects than similar species with green involucral bracts, such as R. pubera. We compared the floral biology of both species and the number of visits to sympatric populations of each species, associating them with the availability of pollen and the pollination system. We verified that species with white involucral bracts are preferred, because there were more visits to R. ciliata in the first 2 h the flowers were open. The peak visitation in R. pubera was 2 h after the flowers opened, when the pollen of R. ciliata was exhausted. Although the involucral bracts of R. pubera are green, the spikelet scales and anthers are white and reflect ultraviolet light. Overall, flowers of R. pubera exhibit fewer white or reflective surfaces and are probably less conspicuous to a bee than those of R. ciliata. It is possible that R. pubera is a second option for visitors after the first 2 h of anthesis. The two different peaks in visitation minimize interspecific competition for pollinators, suggesting that R. ciliata and R. pubera together could attract more generalist pollinators and, instead of competing, facilitate the pollination of both species. Although R. pubera is autogamous and self‐compatible, both wind and insects are important to its reproductive success.     

Nodding Capitula in Superpáramo Asteraceae: An Adaptation to Unpredictable Environment1

Nodding capitula are a striking feature in several Asteraceae species of the Andean superpáramo. In this study, the thermal environment of the nodding inflorescence in Culcitium canescens Humb. & Bonpl. was surveyed by a short-term microclimatic measurement. Temperatures of the nodding inflorescences were higher than temperatures of the surrounding air during the day, while the respective temperatures were comparable during the night. Also, compared to other parts of the plant, the nodding inflorescence provided favorable temperature conditions. An experiment showed that the radiation reflected from the substratum explained most of the variability in the inflorescence temperature. It is suggested that the nodding capitula have evolved to protect the flowers from snow and rain while at the same time ensuring sufficient temperatures for floral development.     

Defense and carnivory: Dual role of bracts inPassiflora foetida

Members of the genusPassiflora are reported to have evolved modifications which kill insects; they have however never been tested for carnivorous syndrome. The flowers ofPassiflora foetida consists of highly reticulate bracts which cover and grow along with the buds and fruits. Removal of bracts from developing bud and fruit resulted in higher predatory damage compared to those where the bracts were intact. These bracts also possess a large number of minute glands which ooze sticky secretion. A variety of tiny insects were found trapped by the secretion of the bracts. The secretion of these glands show high proteases and acid phosphatase activity, two common digestive enzymes found in traps of true carnivorous plants. A high quantity of aminoacids were released from freshly freeze killed ants when incubated in buffer extract of bracts-[¹⁴C] phenylalanine smeared on the glandular surface of bracts was recovered from ovules suggesting potential for absorption of aminoacids. These results suggest a novel role for bracts where primary function is to minimize predatory damage to developing flowers and fruits. The bracts serve as insect traps and also possess the mechanism to digest the trapped insects to obtain free aminoacids.     

Distribution patterns of hummingbird flower mites (Gamasida: Ascidae) in relation to floral availability on Heliconia inflorescences

I analyzed the dispersion patterns of ascid mites (Gamasida:Ascidae) on inflorescences of Heliconia trinidatis in relationto the temporal and spatial distribution of open flowers. Openflowers last only a few hours and are the locations of foodas well as the sole sites from which mites disperse on visitinghummingbirds. All inflorescences were inhabited by populationsof nectarivorous Rhinoseius trinitatis and omnivorous Lasioseiuselegants. All demographic groups of R. trinitatis were associatedstrongly with bracts bearing open flowers on the day of inflorescencecollection or with bracts that would have had an open floweron the day after collection. For L. elegans, only larval distributioncoincided with open flowers. Laboratory observations demonstratedthat mites move throughout an inflorescence at night and preferentiallyaggregate where the next flower will open. Mites enter flowersimmediately at, or just before, anthesis, and unerringly exitopen flowers well in advance of floral abscission. Hummingbirdflower mites exhibit a variety of behavioral adaptations, indicatingextraordinary sensitivity to the physiological and biochemicalnuances of their host inflorescences.     

Johanneshowellia (Polygonaceae: Eriogonoideae), a new genus from the Intermountain West

A new genus ofPolygonaceae subfam.Eriogonoideae is established for what has long been known asEriogonum puberulum. NamedJohanneshowellia in honor of the late John Thomas Howell (1903–1994), a new combination,J. puberula, and a new species,J. crateriorum, are established. The common name Howell's-buckwheat is proposed for the group. The new genus differs from other members of the subfamily in having its involucral structure reduced to a spiral of four to seven bracts each of which is associated with a flower-bearing pedicel and a basal bractlet. In bud and early anthesis the outer two (rarely outer three) bracts may be partially connate, but typically become separate at full anthesis. The connate, ternate, foliar bracts at the base of the node act as a traditionalEriogonum-like involucre surrounding the branches of the inflorescence, the involucral bracts, and the flowers. Members of the new genus are found in the arid Intermountain West of the United States from Inyo Co., California, across Nevada to western Utah.     

Two new endemic species of Ophiorrhiza L. (Rubiaceae: Ophiorrhizeae) from Davao Oriental,Philippines

Two new species of Ophiorrhiza collected from Mt Hamiguitan Range Wildlife Sanctuary in Davao Oriental, Philippines, are herein described and illustrated. Ophiorrhiza erythropilosa is the third Philippine species possessing involucral bracts and is further characterized by its predominantly villose stems that appear red due to strikingly red-violet trichomes, subpersistent bifid stipules, linear involucral bracts without prominent midrib, and 4 mm long urceolate corolla that is villose outside and lightly puberulous inside. Ophiorrhiza hamiguitanensis is characterized by its coriaceous, lanceolate leaves with attenuate apex and base, brochidodromous venation, subpersistent bilobed stipules with slightly recurved acute tips, presence of 1.5–2.5 mm long linear ensiform bracts, heterostylous flowers, clavate calyces and broadly obcordate capsules.     

Multifunctional bracts in the dove tree Davidia involucrata (Nyssaceae: Cornales): rain protection and pollinator attraction

Although there has been much experimental work on floral traits that are under selection from mutualists and antagonists, selection by abiotic environmental factors on flowers has been largely ignored. Here we test whether pollen susceptibility to rain damage could have played a role in the evolution of the reproductive architecture of Davidia involucrata, an endemic in the mountains of western China. Flowers in this tree species lack a perianth and are arranged in capitula surrounded by large (up to 10 cm x 5 cm) bracts that at anthesis turn from green to white, losing their photosynthetic capability. Flowers are nectarless, and pollen grains are presented on the recurved anther walls for 5-7 days. Flower visitors, and likely pollinators, were mainly pollen-collecting bees from the genera Apis, Xylocopa, Halictus, and Lasioglossum. Capitula with natural or white paper bracts attracted significantly more bees per hour than capitula that had their bracts removed or replaced by green paper. Experimental immersion of pollen grains in water resulted in rapid loss of viability, and capitula with bracts lost less pollen to rain than did capitula that had their bracts removed, suggesting that the bracts protect the pollen from rain damage as well as attracting pollinators.     

Floral biology of Nuphar pumila (Timm) DC. (Nymphaeaceae) in China

Anthesis in individual flowers of Nuphar pumila (Timm) DC. occurs for four consecutive days. First-day flowers are protogynous and functionally female. Flowers can open completely on the first day of anthesis. This contrasts with all previous reports, which state that first-day flowers of Nuphar are characterized by partial expansion of the calyx, leaving a distal small triangular opening just above the stigmatic disc. Flowers close completely on the first night of anthesis and remain partially open on the subsequent three nights. During the entire anthesis period the stigmas emit a sweet, fruity odor and the petal nectaries produce visible nectar drops. The stigma of N. pumila is secretory and unicellular-papillate. Pollen grains are monosulcate with long spines. Our observations on the mating system of N. pumila indicate that neither asexual seed production nor spontaneous self-pollination occurs. Cross-pollination of second-, third- and fourth-day flowers produced few seeds. Flowers of N. pumila were mainly pollinated by sweat bees, with flies playing a minor pollination role. No beetle visits were observed. Our insect-pollination observations substantiate the view that the relative contribution of flies, bees, and beetles to pollination in a single Nuphar population depends on two factors: the relative abundance of the insects, and presence of alternative food sources.     

A nodding capitulum enhances the reproductive success of Cremanthodium campanulatum (Asteraceae) at high elevations in the Sino–Himalayan Mountains

Background: Flower orientation is considered an evolutionary response to pollinators and abiotic factors. Several members of the genus Cremanthodium (Asteraceae) growing in alpine habitats in the Sino–Himalayas have conspicuously nodding capitula, the function of which is not understood.

Aims: We investigated the influences of nodding capitula on floral thermal conditions, pollinator visitation rate, pollen viability and achene production in Cremanthodium campanulatum.

Results: (1) the nodding capitulum did not modify internal thermal conditions, but the elongated involucral bracts could shelter pollen from UV-B radiation; (2) water and UV-B radiation seriously reduced the viability of pollen grains; (3) capitula artificially held erect set significantly fewer achenes than nodding ones; and (4) no pollinator preference was observed between capitula that were artificially held erect and natural nodding ones.

Conclusions: This study indicated that the nodding capitulum of C. campanulatum may protect pollen from being damaged by UV-B radiation, being washed away from the anther and stigma, and enable it to avoid rain damage during the wet monsoonal period. Our results suggest that nodding capitula in C. campanulatum possibly confer a selective advantage and may be driven by non-biological agents rather than pollinator attractors.     

全缘叶绿绒蒿的花内热量来源和温度调节功能

为探究全缘叶绿绒蒿( Meconopsis integrifolia )的花内热量来源和温度调节功能,该研究选择在全缘叶绿绒蒿的巴朗山居群,对其进行遮阴及去瓣处理,并采用红外热像仪监测全缘叶绿绒蒿的花内微环境温度日变化及花器官温度,用环境温度计监测环境温度。结果表明:(1)太阳照射显著提高全缘叶绿绒蒿花内微环境温度和花器官温度,全缘叶绿绒蒿的热量主要来源于太阳辐射。花内微环境昼夜温差显著低于环境昼夜温差,全缘叶绿绒蒿的花具有温度调节功能。(2)白天环境温度较高时,太阳照射显著提高全缘叶绿绒蒿花内微环境温度,花瓣会降低花内微环境温度; 夜间环境温度较低时,花瓣闭合会提高花内微环境温度; 花瓣闭合运动降低了花内微环境昼夜温差,产生了保温效果。(3)在太阳照射下,花器官温度差异显著,雌雄蕊温度显著高于花瓣温度,且花器官温度由雌蕊柱头中心点向外递减,全缘叶绿绒蒿能有效调控花器官各部位的温度。综上认为,全缘叶绿绒蒿的花内热量来源于太阳辐射,主要通过花瓣闭合运动降低花内微环境昼夜温差并能在太阳照射下调节各花器官的温度实现温度调节功能。     

重庆赤车属一新种和楼梯草属二新种

描述了自重庆市江津区四面山发现的荨麻科赤车属一新种和楼梯草属二新种：（1）四面山赤车,与融安赤车近缘,区别在于新种的叶的二级脉较少,2枚雌花被片有短角状突起,以及长圆形瘦果;（2）稀柱毛楼梯草,可能与托叶楼梯草近缘,区别在于新种的叶较小,雌总苞苞片,以及雄、雌小苞片均在背面具龙骨状突起,柱头由只3~4根毛组成;（3）四面山楼梯草,与长尖楼梯草近缘,区别在于新种茎的顶端被短柔毛,雄头状花序的花序梗较短,以及雄总苞苞片呈条状船形,背面具龙骨状突起。     

Flower opening and closing ofOxalis martiana

Flowers ofOxalis martiana open in the morning and close in the afternoon repeatedly for 3–5 days in May–July under natural conditions. Both in light and in darkness, the closed flowers opened in reponse to a rise in temperature (thermonasty), but not under the constant temperatures. Transfer from darkness to light along with temperature rise caused rapid flower opening, and at 20 C or higher temperatures, exposure to light caused flower opening even without changing temperature (photonasty). Therefore, the temperature of the night before opening is critical in determining whether the flower opening under natural conditions depends on thermonasty or photonasty. The opened flowers closed about 8–11 hr after the beginning of opening both under natural conditions and constant light-temperature conditions, which suggests that the time of flower closing is determined by endogenous factors. Length of the perianth increased greatly during opening and slightly during closing. Application of actionomycin D or cycloheximide inhibited both the flower opening and closing, probably by suppressing the perianth growth.     

Goodmania (Polygonaceae), a new genus from California

A new genus,Goodmania, is described for the species currently known asOxytheca luteola Parry of central and southern California. A member of Polygonaceae, subfamily Eriogonoideae,Goodmania is an isolated genus, distinct fromEriogonum by the awned bracts, and fromOxytheca by its involucral bracts not forming a distinct tube, yellow flowers, non-glandular pubescence, and small, paired cauline leaves.Goodmania is seemingly related to the Death Valley endemic,Gilmania, but differs in having a whorl of five involucral bracts subtending the cluster of short-pedicellate flowers.     

南溪楼梯草,云南荨麻科一新种

该文描述了自云南东南部发现的荨麻科楼梯草属一新种,南溪楼梯草(Elatostema nanxiense)。此新种与田林楼梯草(E. tianlinense)相似,但其叶具三出脉,边缘具浅圆齿或小钝齿,雄总苞苞片较少,8枚,排成一层,背面有1～3条纵肋,只1枚在顶端具角状突起而与后者相区别。此外,此新种与盘托楼梯草系的广布种盘托楼梯草(E. dissectum)的区别在于其茎被糙伏毛,叶呈椭圆形,具三出脉,雄头状花序的花序梗较短,雄总苞苞片较少,呈宽卵形或横长方形,背面有1～3条纵肋,雄小苞片有缘毛。     

栽培细叶百合的传粉生态

对栽培的细叶百合进行传粉生态学的初步观察结果显示：细叶百合2:30～3:00开始开花,5:30～7:00完全开放;单花开放时间3～4 d,群体花期为12 d。花蜜分泌在开花当日,分泌高峰在9:00～11:00,花蜜分泌与昆虫访花谐调一致。花蜜成分主要为葡萄糖、阿拉伯糖和β－D吡喃果糖,花朵挥发物的成分是软脂酸、软脂酸甲酯、肉豆蔻酸、月桂酸、3－叔戊基戊酮、十三烷和1－甲基,2异丙烯基环丁基乙醇。访花昆虫主要是蝶类、蜂类和蝇类;传粉昆虫为绿豹蛱蝶、大红蛱蝶、绢粉蝶及单齿切叶蜂、小齿淡脉隧蜂,其中小齿淡脉隧蜂的传粉活动最为活跃。晴天时小齿淡脉隧蜂访花频率的日变化与光照的日变化呈极显著的正相关,与温度的日变化呈显著的正相关,与湿度的日变化呈显著的负相关;阴天时与空气温度相关极显著,与空气湿度相关显著。细叶百合以异花虫媒传粉为主,部分自交可育。     

Floral development and evolution of capitulum structure in Anacyclus (Anthemideae,Asteraceae)

Background and Aims

Most of the diversity in the pseudanthia of Asteraceae is based on the differential symmetry and sexuality of its flowers. In Anacyclus, where there are (1) homogamous capitula, with bisexual, mainly actinomorphic and pentamerous flowers; and (2) heterogamous capitula, with peripheral zygomorphic, trimerous and long-/short-rayed female flowers, the floral ontogeny was investigated to infer their origin.

Methods

Floral morphology and ontogeny were studied using scanning electron microscope and light microscope techniques

Key Results

Disc flowers, subtended by paleae, initiate acropetally. Perianth and androecium initiation is unidirectional/simultaneous. Late zygomorphy occurs by enlargement of the adaxial perianth lobes. In contrast, ray flowers, subtended by involucral bracts, initiate after the proximal disc buds, breaking the inflorescence acropetal pattern. Early zygomorphy is manifested through the fusion of the lateral and abaxial perianth lobes and the arrest of the adaxials. We report atypical phenotypes with peripheral ‘trumpet’ flowers from natural populations. The peripheral ‘trumpet’ buds initiate after disc flowers, but maintain an actinomorphic perianth. All phenotypes are compared and interpreted in the context of alternative scenarios for the origin of the capitulum and the perianth identity.

Conclusions

Homogamous inflorescences display a uniform floral morphology and development, whereas the peripheral buds in heterogamous capitula display remarkable plasticity. Disc and ray flowers follow different floral developmental pathways. Peripheral zygomorphic flowers initiate after the proximal actinomorphic disc flowers, behaving as lateral independent units of the pseudanthial disc from inception. The perianth and the androecium are the most variable whorls across the different types of flowers, but their changes are not correlated. Lack of homology between hypanthial appendages and a calyx, and the perianth double-sided structure are discussed for Anacyclus together with potential causes of its ray flower plasticity.     

Comparing Variational Properties of Homologous Floral and Vegetative Characters in Dalechampia scandens: Testing the Berg Hypothesis

The Berg hypothesis posits that, in plants with specialized pollination systems, floral characters should evolve to become integrated with each other and decoupled from vegetative characters. We test this hypothesis by comparing serially homologous and morphologically similar characters in leaves and involucral bracts in the Neotropical vine Dalechampia scandens, which has a specialized pollination system based on resin-collecting bees. The involucral bracts serve a number of specialized floral functions, including signaling and protection, that may put them under stronger selection for precision than the less specialized leaves. The homology and morphological similarity of the leaves and bracts allow us to make a sharper test of Berg’s hypothesis than is possible in most other systems. We found support for the hypothesis in that the bracts had lower coefficients of variation than the leaves for comparable traits. Also in support of the hypothesis, we found essentially zero phenotypic correlations between bracts and leaves at the same time that we found moderate correlations between different leaves and between different bracts. In contradiction to the hypothesis, however, we did not find higher correlations among traits within bracts than within leaves, and we found no evidence of bracts being more developmentally stable than the leaves.     

Biological characteristics of an invasive south African species

Senecio inaequidens, a south African species that has recently invaded parts of Europe, especially human disturbed habitats, was examined under laboratory and glasshouse conditions to assess the germination of achenes, soil seed banks, growth of seedlings and its breeding system. These observations allowed identification of ecological characteristics that may contribute to the invasiveness of the species. Achenes derived from three populations of S. inaequidens were germinated over a wide temperature range (from 14/6 °C day/night temperatures to a constant temperature of 30 °C). They exhibited a rapid achene germination especially when temperatures reached 20 °C. Germination of seeds from soil samples also exhibited a prolonged germination over time suggesting therefore different types of dormancy. Growth response was studied along a nitrogen gradient from 0.33 to 16.0 mmol l^–1 and data were interpreted using growth analysis. Seedlings of S. inaequidens exhibited a slow growth in low nutrient availability and a rapid growth rate when exposed to high nutrient availability, which contributed to producing a large leaf area and biomass. Four types of pollination were studied in S. inaequidens. Hand-pollinations showed that S. inaequidens was mostly self-incompatible. Self-pollinated capitula only showed 7.6% viable achenes, significantly lower than insect-pollinated and cross-pollinated capitula. Some individuals were more self-fertile than others ensuring the production of some offspring in a situation of colonisation. Natural pollination occurred with generalist insects. Most of these results confirm the invasive ability of S. inaequidens.     

A new species of Inga (Fabaceae,Mimosoideae, Ingeae) from Atlantic Forest in Bahia,Brazil

Inga unica Barneby & J. W. Grimes, notable for a syndrome of: a) softly pilose foliage, b) emarginate leafstalks lacking nectaries, c) leaflets 2–3 pairs, of moderate size, d) filiform pliant peduncles, e) globose capitula, f) elongate linear floral bracts persistent after fall of the flowers, g) calyx more than half as long as corolla, and h) thin-textured fruit shattering after falling, is described, figured, and discussed. It appears related toI. (Leptinga) nutans (Vellozo) Martius ex Bentham.