Evidence for rodent pollination in Erica hanekomii (Ericaceae)

Pollination by rodents, originally discovered in Cape Proteaceae, is known from a handful of plant lineages and may be more widespread than current data suggest. Here, we demonstrate the occurrence of this mode of pollination in Erica, the largest plant genus in the Cape flora of South Africa. The localized endemic Erica hanekomii appears to be adapted for pollination by rodents, on account of its low, mat‐forming habit with many dull‐coloured, pendulous flower heads, large volumes (up to c. 30 µL per inflorescence) of sucrose‐dominated nectar, hook‐shaped styles and late winter flowering phenology. Rodents trapped in the vicinity of E. hanekomii plants had many thousands of Erica tetrads in their droppings. Nocturnal laboratory observation showed that Acomys subspinosus (Cape spiny mouse) is adept at manipulating E. hanekomii inflorescences and lapping nectar without destroying gynoecia. During terrarium experiments, pollen was deposited mainly on the rostrum, but also on the chest area and forelegs of A. subspinosus. No birds or insects were seen to visit E. hanekomii in the field. Whereas natural fruit set was 73%, inflorescences from which rodents, but not insects, were excluded using mesh cages, had fruit set of just 3.3%. Exclusion of all visitors resulted in zero seed production. Broken anther rings (a sign of flower visitation in Erica) were observed in 90.1% of flowers in the field and 92% of flowers exposed to rodents in the laboratory, but in virtually none of the flowers from which rodents were selectively excluded. This first record of a non‐flying mammal pollination syndrome in Ericaceae raises interesting new questions about the origins of the rodent pollination system and the selective factors that lead to its evolution. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 163–170.     

The pollination spectrum in the southwestern American cactus flora

The cacti of the American Southwest, defined as the region from southern California to Texas, are surveyed for types of pollination systems and their frequencies. Four types of pollination systems are known to occur in the southwestern cactus flora: bee, hummingbird, hawkmoth, and bat pollination. Two other modes are suspected but not documented: miscellaneous smallinsect pollination and autogamy.—Bee flowers comprise a wide and nearly continuous series of size classes from very small to very large. The large bee flowers, with perianths 5.5 to 12.5 cm in diameter, form a prominent but arbitrarily delimited subcategory in the bee pollination system. Promiscuous flowers and autogamous flowers, if they occur, are included with the medium-sized and small bee flowers in our present classification, due to lack of information, and will have to be separated out when our knowledge is more complete. The overwhelming majority of species in the southwestern cactus flora are bee flowers. Between 39 and 44% of the species are large bee flowers. Another 50 to 56% of the species are classified as medium-sized and small bee flowers; while some of these may turn out to be promiscuous flowers and autogamous flowers, most of them are undoubtedly bee flowers.—Hummingbird, hawkmoth, and bat pollination are conspicuous but statistically minor components of the pollination spectrum, occurring in only one or a few species (see Table 2). Hummingbird pollination has arisen from bee pollination in the Southwest. The species or species groups with hawkmoth and bat pollination, on the other hand, are basically tropical groups which developed their advanced pollination systems in the tropics.Pollination of North American Cacti, V. See alsoGrant & Grant (1979a),Grant & al. (1979),Grant & Grant (1979b),Grant &Hurd (1979).     

Ant pollination of the palaeoendemic dioeciousBorderea pyrenaica (Dioscoreaceae)

The importance of ants for pollination in the dioeciousBorderea pyrenaica (Dioscoreaceae), a Tertiary palaeoendemic plant of the Pyrenees (NE Iberian Peninsula) was studied. The frequency of different visitors (ants, lady beetles, andDiptera) to staminate and pistillate flowers was quantified, and their effectiveness as pollinators was examined by means of fruit and seed set in selective experimental exclusions. Although ants were less abundant on flowers than other visitors, they were the most effective pollinators. Some qualitative factors of this mutualistic ant-plant interaction may account for their effectiveness: the small size of the ants, their high visitation rate to pistillate flowers, and the lack of reduction in viability of the pollen transported on the integument. In addition, the sedentary nature of ants assures their presence during the flowering period. The most abundant floral visitors ofB. pyrenaica were therefore not the most effective pollinators.     

Nectar, floral morphology and pollination syndrome in Loasaceae subfam. Loasoideae (Cornales)

BACKGROUND AND AIMS: Loasaceae subfam. Loasoideae are mostly distributed in South America (sea level to over 4500 m) with a wide range of animals documented as pollinators. The aim was to investigate correlations between nectar parameters, flower morphology, pollination syndrome and phylogeny. METHODS: Nectar was collected from 29 species from seven genera in the subfamily. Concentration and volumes were measured and the amount of sugar calculated. Correlations of nectar data were plotted on a ternary graph and nectar characteristics compared with flower visitors, floral morphology and phylogenetic data. KEY RESULTS: Sugar concentrations are generally higher than reported for most plant families in the literature. The species investigated can be roughly grouped as follows. Group I: plants with approx. 1.5(-3.5) microL nectar with (40-)60-80% sugar and 0.19-2 mg sugar flower-1; with small, white, star-shaped corollas, pollinated by short-tongued bees. Groups II, III and IV: plants with mostly orange, balloon-, saucer-, bowl- or bell-shaped corollas. Group II: plants with approx. 9-14 microL nectar with 40-60% sugar and 4-10 mg sugar flower-1; mostly visited by long-tongued bees and/or hummingbirds. Group III: plants with 40-100 microL nectar with 30-40% sugar and 14-36 mg sugar flower-1, mostly visited by hummingbirds. Group IV: geoflorous plants with 80-90 microL with 10-15% sugar and 8.5-12 mg sugar flower-1, presumably visited by small mammals. Groups II and III include species visited by bees and/or hummingbirds. CONCLUSIONS: Pollinator switches from short-tongued bees via long-tongued bees to hummingbirds appear to have taken place repeatedly in the genera Nasa, Loasa and Caiophora. Changes in nectar amount and concentration appear to evolve rapidly with little phylogenetic constraint.     

Insect-induced wind pollination of the palm Chamaedorea pinnatifrons and pollination in the related Wendlandiella sp.

In Peruvian Amazonia pollination ecology of two palms, Chamaedorea pinnatifrons (Jacq.) Oerst. and Wendlandiella sp. was studied from October 1988 to January 1990. Both palms are dioecious understory species that form locally dense populations.In C. pinnatifrons both sexes flower synchronously during the dry season. Prior to anthesis, the pendulous male inflorescence is inhabited by numerous thrips (Thysanoptera) and Ptiliidae (Coleoptera). Staminate flowers open by a small basal slit between the petals. At anthesis pollen is shed and the movements of the insects inside the flowers trigger pollen release in small clouds. Thus, the powdery pollen becomes airborne and finally air currents act as a vector, carrying pollen to the inconspicuous female plants, which usually are not visited by insects. The term insect induced wind pollination is suggested for this pollination mode. Wendlandiella flowers during four months in the dry season. Male and female plants were not visited by insects. The dry condition of the pollen indicates that anemophily is the pollination mode in Wendlandiella. Fructification is rare but the plants show intense vegetative propagation.The significance of anemophily in the tropical lowland rain forest is discussed.     

Honeyeater (Meliphagidae) pollination and the floral biology of PolynesianHeliconia (Heliconiaceae)

The primary pollinator of Polynesian heliconias,Heliconia laufao andH. paka, is the Wattled Honeyeater,Foulehaio carunculata. This report is the first documentation of pollination by honeyeaters in the genusHeliconia and the first record ofF. carunculata as a pollinator of a plant species. The Polynesian heliconias bear inflorescences that produce 2–4 hermaphroditic flowers per day for a period of 2–3 months. Each flower secretes abundant nectar (125–184 l) with low sugar concentration (15–18% sucrose-equivalents, weight per weight basis) which is available at anthesis just before dawn. Ninety percent of flower visits occur between anthesis and mid-morning. The honeyeaters perch on inflorescence bracts, and probing of the flower results in pollen deposition on the head and bill from where pollen is transferred between flowers. No statements on compatibility can be made forHeliconia paka; however,Heliconia laufao appears to be self-compatible. Calculations of energetic values of nectar of the Polynesian heliconias and Daily Energy Expenditures ofF. carunculata suggest that populations ofH. laufao andH. paka serve as rich energy resources for their pollinators.     

Evolution of a non-flying mammal-dependent pollination system in Asian Mucuna (Fabaceae)

Pollinator shifts are often related to speciation in angiosperms, and the relationship between them has been discussed in several plant taxa. Although limited information on plants pollinated by non-flying mammals in Central and South America and Africa is available, related research has not been conducted in Asia. Herein, I summarize the available knowledge of pollination in Asian Mucuna (Fabaceae), a genus mainly distributed in the tropics, and discuss the evolution of plants pollinated by non-flying mammals in Asia. Nineteen pollinator species have been recorded and pollination systems have been categorized into four types. An examination of the relationship between Mucuna species and their pollinators from the lineage perspective revealed that all species in Mucuna, subgenus Macrocarpa, which are distributed in Asia, are pollinated exclusively by non-flying mammals. Additionally, plants pollinated by non-flying mammals were found to have diverged from bat-pollinated and non-flying mammal-pollinated plants, while plants pollinated by non-flying mammals have evolved multiple times. This is a unique example of evolutionary transition. I hypothesize that the diversification of squirrel species in tropical Asia may have led to the speciation and diversification of Mucuna in Asia. Furthermore, the behavioural and ecological characteristics of bats and birds in Asia differ from the characteristics of those in other regions, implying that Asian Mucuna species do not rely on bat or bird pollinators. The adaptation of floral characteristics to pollinators is not well understood in Asia. Mammal-pollinated plants in Asia may have evolved differently from those in other regions and have unique pollination systems.     

Hydrophilous pollination and reproductive morphology in the seagrassPhyllospadix scouleri (Zosteraceae)

In the seagrassPhyllospadix scouleri (Zosteraceae) floral aggregates (spadices) occur on short lateral axes produced by subordinate vegetative shoots. The filamentous pollen is both dispersed on the surface of the sea as well as below the surface. In surface pollination, snowflake-like search vehicles (pollen rafts) float and collide with any rigid female inflorescences that emerge from the water surface. In submarine pollination, collinear bundles of pollen grains are dispersed. Analysis of seed set indicates ovule position within the inflorescence to affect likelihood of fertilization in submarine, but not surface-pollinated inflorescences. Agamospermy appears to be unlikely, but the strongly female-biased shoot sex ratio remains to be explained.     

Observations on pollination inSarcostemma (Asclepiadaceae)

Field observations on pollination in New World species of the genusSarcostemma R. Br. sensuHolm are reported. Morphological and anatomical comparisons of pollinated flowers ofSarcostemma subg.Ceramanthus Kuntze (New World) andSarcostemma subg.Sarcostemma (Old World) are presented.     

The pollination ecology ofBletilla striata (Orchidaceae)

Anthecological observations of an orchidBletilla striata were carried out in Kobe, Japan. The purplish pink flowers were visited by 26 species of insects belonging to Hymenoptera, Diptera and Lepidoptera, although the flower did not offer food rewards of any kind. Candidates for pollination were seven species of aculeate Hymenoptera with a body size that well fitted the space in between the column and labellum. The most effective candidate was the male of the longhorn beeTetralonia nipponensis, judging from its abundance and legitimate intrafloral behavior. The female ofT. nipponensis and the other six species also contributed to promote out-crossing, but to a lesser extent than the maleT. nipponensis, since they occasionally exhibited illegitimate intrafloral behaviors for pollination and also, their flower visitation rate was lower.     

Size-dependent pollination efficiency in Anchusa officinalis (Boraginaceae): causes and consequences

Summary Bumblebees foraging on the self-incompatible Anchusa officinalis fly between near neighbour plants and between near neighbour inflorescences within plants. Although many-flowered plants attracted most bumblebees these plants received fewer visits on a per flower basis than smaller plants, and each bumblebee visited a smaller proportion of the flowers. The calculated effective visitation rate per flower was highest on plants of an intermediate size. If pollen-carryover was assumed to be limited the most efficient plant was predicted to be smaller since the proportion of fertilized flowers per bumblebee visit is expected to decrease further on the largest plants in relation to the total flower number. These predictions were tested by measuring fruit-set in the field. The percentage fruit-set decreased with plant size at all sizes that were investigated. That the most efficient plant was small indicates that pollen-carryover was indeed limited. However, the low percentage fruit-set associated with large size did not present a serious problem since the total estimated seed production per plant still increased with size. Selection favoring smaller plants may be low or absent in Anchusa.     

The remarkable weevil pollination of the neotropicalCarludovicoideae (Cyclanthaceae)

Further evidence for a specialized plant-pollinator relationship in the weevil pollinatedCarludovicoideae (Cyclanthaceae) is presented, with emphasis on the previously uninvestigated genusSphaeradenia. Weevils use the carludovicoid inflorescence for feeding, shelter, mating, and oviposition, the last-mentioned activity in the usually caducous staminate flowers. While visiting the inflorescence, they effect pollination. Adaptations, advantages, and consequences for this sophisticated system, both of the plants and pollinators, are discussed.     

Oriole pollination ofErythrina breviflora (Leguminosae): Evidence for a polytypic view of ornithophily

Erythrina breviflora is visited by large numbers of passerine birds of which orioles (Icterus: Icteridae) are the primary pollinators. The flowers produce large quantities of nectar but they are rarely visited by hummingbirds. Inflorescence and floral morphology, and low levels of sucrose in the nectar probably explain the rarity of foraging hummingbirds. A comparison of Old WorldErythrina and their pollinators with New World species pollinated by orioles and hummingbirds suggests that parallel evolution has occurred. When the comparison is expanded to include other species pollinated by orioles, it is clear that various New WorldIcteridae, Thraupidae, etc. are ecological equivalents of Old WorldOriolidae, Pycnonotidae, Sturnidae, etc. and that flowers pollinated by these birds have similar characteristics.     

Floral scent chemistry and pollination ecology in phytelephantoid palms (Arecaceae)

The subfamilyPhytelephantoideae comprises three genera (Ammandra, Aphandra, andPhytelephas) and seven species of dioecious palms. The floral scents ofAmmandra dasyneura, A. decasperma, Aphandra natalia, Phytelephas aequatorialis, P. macrocarpa, andP. seemannii were analyzed by gas chromatography-mass spectrometry. We studied the pollination biology ofA. natalia, P. aequatorialis, andP. macrocarpa, and tested how the synthetically produced main constituents of the floral scents ofAphandra andPhytelephas attracted insects in two natural populations ofPhytelephas. The genera are distinct in terms of floral scents.Ammandra has sesquiterpenes,Aphandra (+)-2-methoxy-3-sec-butylpyrazine, andPhytelephas p-methyl anisol. These constituents dominated the scents quantitatively and qualitatively. The similarity between scents of male and female inflorescences was 76.5% inAmmandra, 84.2% inAphandra, and >99% inPhytelephas. Different species ofAleocharinae (Staphylinidae) pollinateAphandra natalia andPhytelephas species and reproduce in their male inflorescences.Derelomini (Curculinoidae) andMystrops (Nitidulidae) only visit and pollinatePhytelephas in which male inflorescences they reproduce. A species ofBaridinae (Curculionidae) only visits and pollinatesAphandra natalia, and reproduces in its female inflorescence. The apparent reliance on one or a few floral scent constituents as attractants and few and specific pollinators may indicate co-evolution. Sympatric species ofPhytelephantoideae have different scents. We suggest that species with similar scents have allopatric distributions due to the absence of a pollinator isolation mechanism.     

濒危植物丰都车前的繁育系统与传粉生物学研究

丰都车前为三峡库区特有珍稀濒危植物,目前仅在重庆市江津支坪镇涂家村附近大中坝上有一小种群分布。以野生种群和移栽种群为对象,通过野外观察和人工授粉实验对丰都车前传粉生物学及繁育系统进行研究。结果表明:(1)丰都车前单花花期约4 d,花序花期约8 d,种群花期约3个月;单花开花进程可分为6个时期:花蕾期、雌蕊伸长期、雄蕊初露期、花瓣展开期、花瓣反折期、枯萎凋谢期。(2)花瓣展开前,柱头先伸出花冠,达最高可受性,之后花瓣展开雄蕊才露出,而此时大部分柱头已枯萎失活丧失可受性,只有少部分具可受性。表现出雌雄异熟的特征,一定程度避免了自交的发生。(3)花粉活力始于雄蕊初露期,花瓣展开、花药开裂时花粉活力最高,至枯萎凋谢前均具有活力。花粉活力最高期避开了柱头可受性最佳期,仅有少部分重叠。(4)繁育系统检测显示,花粉/胚珠比为15373.29,杂交指数为3。结合套袋实验表明,丰都车前可能同时具有风媒和虫媒传粉机制,传粉昆虫为中华蜜蜂、食蚜蝇,但数量偏少,不存在无融合生殖,异交为主,部分自交亲和,有时需要传粉者。(5)开花时期避开了其分布地洪水淹没期,能够顺利完成开花授粉及座果。综上所述,丰都车前的花部特征、开花物候和繁育系统为适应分布地夏季洪水水淹胁迫提供了一定的生殖保障,促进了该种群的续存。     

Beetle pollination of peacock moraeas (Iridaceae) in South Africa

The peacock moraeas are South African geophytes withIris-like flowers that are characterized by broad outer tepals with iridescent spots. Three of the seven species were studied and found to be exclusively visited and pollinated by beetles. The primary pollinators were hopliine (Scarabaeidae) beetles in the generaPeritrichia, Monochelus, Anisonyx andAnisochelus. These beetles visit flowers for nectar, pollen and mating.Peritrichia rufotibialis was the primary pollinator ofMoraea villosa, P. abdominalis was the primary pollinator ofM. tulbaghensis, whileMonochelus sp. nov. was the primary pollinator of the third species,M. neopavonia. High levels of fruit set attest to the effectiveness of these beetles as pollinators. Average fruit set ranged from 83.5 ± 34.8% (M. villosa) to 97.2 ± 11.6% (M. tulbaghensis). Average seed set in the various populations was more variable, but was also quite high, ranging from 77.3 ± 40.8 (M. villosa) to 342.2 ± 115.0 (M. neopavonia) seeds per capsule. Although hopliine beetles are abundant and diverse in southern Africa and commonly visit flowers, this is the first well-documented report of pollination by these beetles.     

Breeding systems and pollination inVigna minima (Leguminosae,Papilionoideae)

Four types of floral breeding systems—(i) chasmogamy, (ii) aerial pseudocleistogamy, (iii) subterranean pseudocleistogamy and (iv) obligate subterranean true cleistogamy—are observed in the populations ofVigna minima inhabiting the Western Ghats (India). Five categories of phenotypes are recognized based on the number and kinds of floral breeding systems found in a given individual. The frequencies of different categories of phenotypes not only show intra- and interpopulation variation, but also fluctuate from generation to generation suggesting differences in the genetic structure of populations. This polymorphism in the breeding system of a single species is unique and may be adaptive. Obligate subterranean true cleistogamy and amphicarpy appear to be adaptations to jungle fires and soil erosion.—The flowers are of the flag-blossom type and insect visitors act as tripping agents. The tripping mechanism together with the polymorphic floral breeding system result in a balanced mixture of selfing and outcrossing. Such a recombination system may enhance the fitness ofV. minima which is essentially a colonizing species.     

外来入侵植物意大利苍耳的传粉生态学特性

从开花物候、花粉活力、柱头可受性、传粉媒介、花粉散布距离、雄性、雌性功能以及繁育系统等方面系统地研究了意大利苍耳的传粉生态学特性,旨在为揭示该物种成功入侵的机理提供科学依据。研究结果显示:意大利苍耳种群的花期较长,雌花花期可达40d,雄花花期可达30d。其雄性和雌性功能都很强,意大利苍耳单株雄花序和花粉量分别高达3847个/株和37903037粒/株,单株雌花序和单株胚珠数高达3847个/株和7694枚/株,种群平均结实率高达80.59%,即平均每株个体能形成6200枚成熟的果实。花粉活力的日变化呈单峰型曲线,早晨和傍晚的花粉活力最低,14:00时的活力高达99%。雌蕊柱头一经伸出总苞即具有可受性,可受期长达7d,开花后的3—4d柱头的可受性最强。意大利苍耳为风媒传粉植物,在4.5km/h的微风条件下,其花粉的散布距离可达45m,大量的花粉集中分布在距花粉源0—20m的范围内,这对于往往高密度连片分布的意大利苍耳种群来说无疑是一种高效的传粉策略。意大利苍耳的交配系统灵活多样,盛花期自然结实率达到100%,套袋试验结果表明该植物自交亲和,自株自然授粉的结实率高达93%。表明较长的花期、大量的雌雄花序及花粉数量、较高的花粉活力、较长的柱头可授期、较远的花粉风媒散布距离、混合交配系统、以及较高的结实率是意大利苍耳繁殖成功的重要保障,也是其成功入侵的重要原因。     

野慈姑(Sagittaria trifolia)人工授粉的有效性:授粉工具与花粉保存

人工授粉作为一项关键性技术在生态研究与农业生产领域广泛应用,但是授粉过程中的工具有效性以及花粉保存尚未受到研究者的关注。以植物野慈姑为试验材料,研究了人工授粉方式和花粉保存条件对繁殖的影响。两年的结果表明,相较于直接接触式授粉,借助毛刷工具授粉显著降低了野慈姑的座果概率和种子数。花粉经过半小时野外高温保存后,其授粉的座果概率显著高于对照组,但单果种子数和种子大小保持稳定。综上,在野慈姑的人工授粉中应避免使用毛刷等工具,离体花粉可置于野外环境适当保存。在相关应用中,研究人员应当警惕工具授粉造成的负面影响,注意比较工具与其他授粉方式的等效性,保证实验的科学性。     

The pollination syndrome ofDeplanchea tetraphylla (Bignoniaceae)

The reproductive structures ofDeplanchea tetraphylla (Bignoniaceae) exhibit a significant number of unusual features: inflorescence with an apical platform; flowers yellow, short-tubed, strongly zygomorphic; mouth closed through lateral compression; stamens and style long-exserted, erect or slightly reclined; nectar dark brown, exposed in the spoon-shaped lowermost corolla lobe and apparently acting also as a visual cue. These features suggest a highly elaborate syndrome for bird pollination: the birds (probably lorikeets) perch on the inflorescence platform and bend downwards to take up the exposed nectar, thus touching the exserted anthers and stigmas with the throat or breast. The likely evolution of this syndrome by additive steps, effecting a change from head up to head down position of the pollinator, can be traced from the floral structure of the remaining four species of the genus.