Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

Density-dependent mating and reproductive assurance in the temperate forest herb Paris quadrifolia (Trilliaceae)

In animal-pollinated plants, autonomous selfing provides reproductive assurance under conditions of infrequent pollinator visits or a lack of mates, but few data are available for wind-pollinated species or species with combined insect and wind-pollination, for which it is often assumed that pollen availability does not limit reproduction. In this study, the capacity of autonomous selfing was investigated in the temperate forest herb Paris quadrifolia, and an emasculation experiment was performed under natural field conditions to investigate the contribution of autonomous selfing to total seed set across a continuous gradient of densities of flowering conspecifics. In the absence of wind or pollinators, autonomous selfing was observed through anthers approaching stigmas at the end of flowering and the capacity for autonomous pollination was about 0.34. Under natural conditions, considerable outcross pollination was observed, but the proportion of ovules successfully fertilized significantly decreased with decreasing density of conspecifics when flowers were emasculated, but not when flowers were left intact. These results indicate that autonomous selfing resulted in reproductive assurance (RA = 0.16) and thus support the hypothesis that autonomous selfing can also provide reproductive assurance in wind-pollinated species.     

珍稀濒危植物单性木兰传粉生态学研究

野外定位观测了单性木兰的花期物候、访花昆虫种类、访花频率、访花行为和同花期植物种类,并对其花粉散播距离和繁育系统进行了检测,旨在探讨制约单性木兰传粉过程的某些因素,为单性木兰的保护生物学提供科学依据。结果表明,单性木兰雄株和雌株在花期物候上存在差异,雌株在结实上存在明显的大小年现象。单性木兰以虫媒传粉为主,雄株访花昆虫30种,雌株访花昆虫仅14种,雌株和雄株共有的访花昆虫仅6种。单性木兰同花期植物共17种,其访花昆虫种类大部分与单性木兰的访花昆虫种类相同。蜂类和蝶类均非单性木兰的传粉者,其结实率低的原因不仅与访花昆虫种类少和访花频率少有关,更与其缺乏有效的传粉昆虫有关。     

Contrasting flowering phenology and species richness in abiotically and biotically pollinated angiosperms

Abstract Biotic pollination is thought to correlate with increased interspecific competition for pollination among plants and a higher speciation rate. In this study we compared patterns of flowering phenology and species richness between abiotically (wind) and biotically pollinated plants, using phylogenetically independent contrasts. We compiled phenological data from eight local seasonal floras, in which we found geographically overlapping sister clades. Of 65 documented origins of wind pollination, we were able to use up to 17 independent contrasts. In contrast to previous studies we found no difference in global species richness between wind- and biotically pollinated sister clades. Regarding phenology, we found wider phenological spread in biotically pollinated clades, earlier flowering onset in wind-pollinated trees, but no difference in duration of flowering between pollination modes. These results corroborate previous views that niche space is more constrained for wind-pollinated species, and that niche partitioning is less important between wind-pollinated plants compared to plants pollinated by animals.     

Variable flowering phenology and pollinator use in a community suggest future phenological mismatch

Recent anthropogenic climate change is strongly associated with average shifts toward earlier seasonal timing of activity (phenology) in temperate-zone species. Shifts in phenology have the potential to alter ecological interactions, to the detriment of one or more interacting species. Recent models predict that detrimental phenological mismatch may increasingly occur between plants and their pollinators. One way to test this prediction is to examine data from ecological communities that experience large annual weather fluctuations. Taking this approach, we analyzed interactions over a four-year period among 132 plant species and 665 pollinating insect species within a Mediterranean community. For each plant species we recorded onset and duration of flowering and number of pollinator species. Flowering onset varied among years, and a year of earlier flowering of a species tended to be a year of fewer species pollinating its flowers. This relationship was attributable principally to early-flowering species, suggesting that shifts toward earlier phenology driven by climate change may reduce pollination services due to phenological mismatch. Earlier flowering onset of a species also was associated with prolonged flowering duration, but it is not certain that this will counterbalance any negative effects of lower pollinator species richness on plant reproductive success. Among plants with different life histories, annuals were more severely affected by flowering–pollinator mismatches than perennials. Specialized plant species (those attracting a smaller number of pollinator species) did not experience disproportionate interannual fluctuations in phenology. Thus they do not appear to be faced with disproportionate fluctuations in pollinator species richness, contrary to the expectation that specialists are at greatest risk of losing mutualistic interactions because of climate change.     

Wind pollination and insect pollination of two temperate willow species,Salix miyabeana and Salix sachalinensis

Pollination systems of Salix miyabeana and Salix sachalinensis were studied at a riverside in northern Japan in order to measure the balance of wind pollination and insect pollination. In 1996, 19 clones of each species were selected, and seed-set success by a net-bagging (to exclude insect visitation) and an artificial pollination (to remove pollen limitation) were compared to by natural pollination. For S. miyabeana, the same experiment was repeated in two populations in 1997. Proportion of seed set through wind pollination dominated in both species in this study. Pollen limitation was common under natural conditions, and S. sachalinensis relied more on insect pollination for seed production than S. miyabeana. Meteorological factors such as precipitation and hours of sunshine during the flowering season influenced the potential reproductive activity of the willow between years. In the wet and cloudy spring of 1996, clones which obtained high seed set depended more on insect pollination for both species, whereas in the dry and sunny spring of 1997, such clones depended more on wind pollination for S. miyabeana. Because the efficiency of wind pollination seemed to be more sensitive to fluctuating weather conditions than insect pollination, insect pollination was considered to play an assurance role for seed production in these willows.     

Pollination Biology of Distylous Rubiaceae in the Atlantic Rain Forest, SE Brazil

Abstract: Data on pollination biology constitute important clues for the comprehension of pollen flow and genetic differentiation in plant populations. Pollinator type, availability and behaviour may modify morphological and mating patterns in populations of typically distylous species. This study investigates the pollination biology of four distylous species of Rubiaceae in the Atlantic rain forest, SE Brazil. Data on flowering phenology, floral lifespan, stigmatic receptivity, pollen availability, nectar volume and concentration, and pollinator activity were collected. The species studied flower sequentially throughout the wet season, and produce terminal inflorescences which bear small, tubular, diurnal, nectariferous flowers. Despite these similarities, some of the species studied are pollinated by different groups of pollinators, probably due to their distribution, availability of flowers and corolla length. On the other hand, pollinator specificity does not seem to be so important for distylous species. Long mouthparts, like those of most of the recorded pollinators, may reach lower sexual organs and, together with the self- and intramorph-incompatibilities observed, be sufficient to perform legitimate pollination and maintain levels of intermorph mating.     

A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernández Islands (Chile)

A survey of the reproductive features of the angiosperm flora of the Juan Fernández Archipelago (Chile) is presented to provide a species-based review of reproduction and pollination, to identify generalizations associated with these systems, to understand the evolution of these features, and to utilize these data to promote conservation. The collection of original data was extensive, based on our own fieldwork, and was combined with data from existing literature. Data recorded include habit, sexual system, flower size, shape, and color, and the hypothesized pollination system of the first colonizers. In addition, the data on compatibility, presence and type of dichogamy, observed floral visitors, presence of floral rewards, and currently known pollination systems are summarized. Pairwise comparisons of different features are tested for statistical association. The flora is typically composed of perennials. The majority of the species have very small or small flowers. Inconspicuous flowers (i.e., a shape character describing flowers with no optical attraction) are widespread, as are dish-shaped flowers. Green is the most frequent flower color, followed by white and yellow. Most species are hermaphroditic, 9% are dioecious, and 9% are monoecious. Some 30% of the species are protandrous and 7% protogynous. Detailed studies of compatibility of about 14% of the flora indicate that 85% of these species are self-compatible (SC). Although most species studied are SC, their level of autogamy is low. Nevertheless, selfing mediated via geitonogamy is the most frequent mechanism of pollen transfer. Outcrossing is mainly achieved through dioecy and self-incompatibility (SI), promoted by dichogamy in the hermaphroditic flowers, and facilitated by wind pollination. About 55% of the species offer nectar rewards, and only 2% offer pollen rewards. Floral visitors are rare to uncommon. Two hummingbird species, one of them endemic, are considered as pollinators for 14 plant species. Flies, moths, and beetles are the native insect visitors to flowers, but they have been documented on only 11 plant species (7%). Even insect visits to these few species were rarely observed. Given the infrequent, irregular, and imprecise nature of native insect association with flowers, there is no certainty that any of the species are truly insect pollinated. Two species of introduced ants and a new endemic bee were recorded as well; however, neither is likely currently important to the pollination of the native flora species. About 9% of the extant flora is currently bird pollinated, and we hypothesize that 47% is wind pollinated. However, we propose that most of the colonizers were ancestrally either insect or wind pollinated. There is association between a number of current floral features and the hypothesized pollination of colonizers. Therefore, to a large extent the flower color, shape, and size of the extant flora may express the pollination syndromes of colonizers rather than representing extant pollination. In addition, the presence of nectar in many species of extant flora does not necessarily indicate biotic pollination. Thus, studies of the reproductive biology on oceanic island plants need to be conducted species by species before broad generalizations can be made, because the observed features can be misleading. Possible changes in the pollination system were assessed by comparison of species for which there are reliable data with the hypothesized pollination of their colonist progenitors. The wind-or bird-pollinated species have retained the pollination system of the colonizers. In other instances, species seem to bear a different pollination system: from ancestral insect systems to current hummingbird-or wind-pollination systems. The lack of alternative means of biotic pollination seems to have led in a number of instances to anemophily—in essence a default pollination system. The lack of strong selection pressure for wind pollination and the relative youth of the archipelago may help explain why the features associated with wind pollination in these species are not so obvious. Because there are many recorded extinctions of vascular plants from islands versus those from continental areas, it is imperative to invest additional effort in protecting the remaining island species. Conservation or restoration programs cannot be effective without a deep and broad understanding of the reproductive biology of the plants. In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The ultimate fate of some species may depend on preserving the plant-hummingbird relationship, including the web of organisms that affect both plant and pollinator. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species.     

The assembly of plants used as nectar sources by hummingbirds in a Cerrado area of Central Brazil

Studies on hummingbird–plant interactions commonly use a pollination approach emphasizing mutualistic relationships. But floral resources are often used opportunistically by these birds. Plant–pollinator assemblies and pollination sustainability will depend both on the well-adapted plants and other potential floral resources. The Cerrado, Neotropical savannas of Central Brazil, has ca. 7.5 % of its flora supposedly adapted to hummingbird pollination. But detailed information about flowers effectively used by hummingbirds at community level is still lacking. Hence, we recorded all plant species visited by hummingbirds, to determine how these nectariferous flowers were distributed in time and space in different plant formations of a Cerrado area, and also the hummingbird species that visit them. The study was conducted between March 2007 and December 2008 in the Panga Ecological Station. Data regarding flowering phenology, floral morphology and visitation were collected monthly. Forty-six nectariferous species from 39 genera and 17 families were recorded, most with annual flowering dynamics and tubular flowers. But only 21 species had a combination of traits fitting classic ornithophilous syndrome. For the remaining species hummingbird visitation was ascertained from observations at the study site or other sites in the region. Eight hummingbird species occurred in the area and were recorded visiting directly 36 plant species. The study area presented a relatively low number of ornithophilous plants, but a great habitat diversity and many non-ornithophilous plants that hummingbirds used as nectar sources. Therefore, in the studied Cerrado, the diversity of environments and nectariferous plants favour the maintenance of resident and migrant hummingbirds.     

First flowering of wind‐pollinated species with the greatest phenological advances in Europe

Increasing risk of pollinosis (hay fever) is one of the most anticipated consequences of climate change on human health. Wind‐pollinated plants are representative of allergenic species because they include species with the highest capability of causing allergy‐related diseases in humans. Therefore, changes in wind‐pollinated species may reflect impacts of climate change on allergenic plants. In particular, flowering is one of the developmental stages most affected by climate change. This report specifically addresses changes in flowering dates that have occurred during the three decades 1971–2000 as a function of pollination mode and woodiness. The assessment is made using a phenological data set comprising trends of flowering dates of 29 species in 983 locations in Europe. Linear mixed models assessing the statistical significance of trends while adjusting for spatial correlation are used. The main results indicate for the first time that the onset of flowering of wind‐pollinated plants advanced more than for insect‐pollinated plants, while full flowering phases tended to advance less. These novel findings are contrary to the results of Fitter and Fitter (2002) for Oxfordshire, who reported larger advances of insect‐pollinated plants. Onset of flowering and full flowering of insect‐pollinated species are more likely to advance for seasons early in the year; instead, wind‐pollinated plants showed no dependence of trends on the season (first flowering) or a decreased advance of phases that are early in the year (full flowering). The effect of woodiness could not be unambiguously defined, but seems to be of minor importance. The presented findings suggest a lengthening of the flowering period in general, which might lead to an increasing time of exposure to airborne pollen of allergic subjects, with consequent likely increment in severity and incidence of allergic symptoms.     

Insect pollinators of anise plants (Pimpinella anisum L.) and the important role of honey bees (apis mellifera L.) on their yield productivity

The present investigation aimed to identify the insect pollinators and the efficiency of honeybees in pollination of anise plant during flowering periods of 2008 and 2009 seasons at Assiut Governorate, Egypt. Obtained results indicated that the total insect counts attained 11 species, belonging to four orders; Hemiptera (one species), Coleoptera (two species), Diptera (five species) and Hymenoptera (three species). Honeybees were the most abundant of hymenopterous insects during anise flowering and the daily peak activities were detected between 12.00 noon and 2.00 p.m. for the two seasons. Regarding the effect of prevailing air temperature and relative humidity percentage (RH%) on the occurrence of insect pollinators, it is clear that the effects of the two factors were moderate in the two seasons. Also, the results showed a significant effect of open pollination on the seed yield/feddan (1024.12 kg) followed by honeybees pollination (781.55 kg), while insect exclusion recorded the lowest value (300.24 kg).     

Distribution and phenology of climbers of Kumaun in Central Himalaya,India

A total of 105 climber species were recorded and studied for their altitudinal distribution and phenology. Relatively higher percentage (54%) of deciduous species was recorded. In general, the climbers were distributed throughout the region between altitudes of 300–3500 m. However, maximum (more than 60%) species were confined either below or at around 1500 m altitude. Species richness declines gradually towards the higher altitudes, the decline was more sharp in evergreen species. The different phenological activities were usually governed by the seasonal variations in a year. Seasonality of flowering suggested the prevalence of insect pollination. Whereas, fruit types indicated equal sharing of biotic and abiotic dispersal. Study concludes, the distribution and nature of various climber species of Kumaun in Indian Central Himalaya, depends largely upon altitude and its climatic variable temperature. Whereas, the seasonality of rains plays an important role in seasonal progression of phenophases.     

华南云实传粉生态学(英文)

在广东鼎湖山,华南云实(LaesalpiniacristaL.)始花期从2月或3月开始,开花后约1周达到盛花期,盛花期持续2~4d,花期可持续到4月底。华南云实基本繁育系统为雌蕊先熟、自交不亲和的异花受精。华南云实传粉方式为风虫媒。其主要传粉者为膜翅目昆虫,该种是云实属植物中首次报道的可行风媒传粉的种类。描述并探讨了华南云实花的结构与其主要传粉者木蜂之间的互动适应,并讨论了传粉生物学实验中的一些处理方法对华南云实开花过程的影响。     

华南云实传粉生态学

在广东鼎湖山,华南云实(Laesalpinia crista L.)始花期从2月或3月开始,开花后约1周达到盛花期,盛花期持续2～4 d,花期可持续到4月底.华南云实基本繁育系统为雌蕊先熟、自交不亲和的异花受精.华南云实传粉方式为风虫媒.其主要传粉者为膜翅目昆虫,该种是云实属植物中首次报道的可行风媒传粉的种类.描述并探讨了华南云实花的结构与其主要传粉者木蜂之间的互动适应,并讨论了传粉生物学实验中的一些处理方法对华南云实开花过程的影响.     

Divergence on floral traits and vertebrate pollinators of two endemic Encholirium bromeliads

Shifts in pollen vectors favour diversification of floral traits, and differences in pollination strategies between congeneric sympatric species can contribute to reproductive isolation. Divergence in flowering phenology and selfing could also reduce interspecific crossing between self‐compatible species. We investigated floral traits and visitation rates of pollinators of two sympatric Encholirium species on rocky outcrops to evaluate whether prior knowledge of floral characters could indicate actual pollinators. Data on flowering phenology, visitation rates and breeding system were used to evaluate reproductive isolation. Flowering phenology overlapped between species, but there were differences in floral characters, nectar volume and concentration. Several hummingbird species visited flowers of both Encholirium spp., but the endemic bat Lonchophylla bokermanni and an unidentified sphingid only visited E. vogelii. Pollination treatments demonstrated that E. heloisae and E. vogelii were partially self‐compatible, with weak pollen limitation to seed set. Herbivores feeding on inflorescences decreased reproductive output of both species, but for E. vogelii the damage was higher. Our results indicate that actual pollinators can be known beforehand through floral traits, in agreement with pollination syndromes stating that a set of floral traits can be associated with the attraction of specific groups of pollinators. Divergence on floral traits and pollinator assemblage indicate that shifts in pollination strategies contribute to reproductive isolation between these Encholirium species, not divergence on flowering phenology or selfing. We suggest that hummingbird pollination might be the ancestral condition in Encholirium and that evolution of bat pollination made a substantial contribution to the diversification of this clade.     

Role of pollination in yield and physicochemical properties of tomatoes (Lycopersicon esculentum)

Very little is known about pollination and its effects on the yield and physicochemical properties of flowering plants in tropical countries. Wind and insect pollinators are among our natural resources because pollination is the most important ecosystem service performed by wind and insects, and is vital to the socio-economic status of human beings. In this experiment, different pollination methods for tomato plants were examined. Self-pollination was encouraged by covering the plants with a plastic sheet. Wind and insects were excluded from these plants, and thus only self-pollination was possible. The experiment occurred during the flowering stage. Wind-pollinated plants were covered with a muslin cloth, which excluded insects, and only wind could pass through the cloth. For insect pollination, plants remained uncovered, allowing free access to insects to pollinate the flowers. At fruit maturity, when fruits were completely red, fruits from each treatment were harvested on the same date and under the same conditions. Results illustrated the substantial importance of insects as pollinators of tomato crops. Open field had greater tomato yield and positive effects on physicochemical properties on fruit than under self and wind pollination.     

Pollination Ecology of Caesalpinia crista （Leguminosae： Caesalpinioideae）

The flowering phenology, pollination ecology and breeding system of Caesa/pinia crista L.were studied in Dinghushan Nature Reserve, Guangdong, China. The species started blooming in Februaryor March, then last till late April. It took about one week from first flower appearance to its full blooming,which lasted for 2-4 d. The pollen-ovule ratio was 18 000~500. The breeding system was self-incompatible,and protogynous xenogamy. Hymenoptera constituted the major group of pollinators. The pollination typeis ambophily, the species could be pollinated by wind if the pollinators were unavailable: this is the firstrecord of ambophily in the genus Caesalpinia. The floral structure adaptation to the pollinating behavior ofcarpenter bees was described. The influences of artificial treatments in pollination biological studies onthe flowering and fruiting of the plants were also discussed.     

Mechanisms that limit pollinator range in Ericaceae

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.     

Breeding system of the subantarctic plant species Pringlea antiscorbutica R. Br. and search for potential insect pollinators in the Kerguelen Islands

The reproductive biology of Pringlea antiscorbutica R. Br. (Brassicaceae), an endemic species from the southern Indian Ocean islands, is investigated here. Controlled crosses were performed between plants grown in a common garden on the Kerguelen Islands. Searching for potential insect pollinators, we investigated the flying ability of all insects known from the Kerguelen Islands. We showed evidence for self-compatibility, low pollen limitation in the absence of a biotic pollen vector and slight selfing depression in P. antiscorbutica. The insects present on the islands are mostly wingless and are not likely to be efficient pollinators of this species. Since P. antiscorbutica shows morphological adaptations to insect pollination despite the absence of pollinators in its present range, we conclude that this species recently evolved from insect pollination to wind and/or autonomous pollination.     

Reproductive Traits and Phenology of Plants in Tropical Dry Evergreen Forest on the Coromandel Coast of India

Qualitative reproductive traits of 84 plant species belonging to 41 families were studied in tropical dry evergreen forest on the Coromandel coast of India. Majority of species had rotate type, white-coloured, scented flowers, rewarding nectar and pollen and pollinated chiefly by bees. An association between floral traits and pollination spectrum is evident. Bee pollination was prevalent in pollination systems. Among the fruit types, drupe and berry were common in black and red colour respectively, and dispersed by zoochorous mode. Seeds of brown- and green-coloured dry fruits, without any reward were disseminated by wind and explosion. The reproductive phenophase of trees and lianas occurred mostly during the dry period from January to June, which receives rainfall of less than 50 mm a month. However, shrubs showed a peak in flowering and fruiting in wet period. Detailed phenological observations of 22 woody species revealed a seasonal and unimodal pattern in flowering. Although some species were in flower round the year, flowering activity was skewed towards the dry season. The fruiting activity showed a bimodal pattern, one peak in dry season and another in wet season. Many species displayed a temporal aggregation in flowering and fruiting. The significant relation was obtained between reproductive traits and phenology of plants in the tropical dry evergreen forest.