Direct and Indirect Effects of Nectar Robbing on the Pollinating Behavior of Patagona gigas (Trochilidae)1

In central Chile, nectar robbing of Puya coerulea (Bromeliaceae) flowers by the austral blackbird, Curaeus curaeus, had a direct impact on the plant via removal of floral resources and, in some cases, damage or even destruction of flowers. These robbing visits also likely had an indirect impact on plant fitness due to the decreased rate of legitimate visits by the hummingbird Patagona gigas to inflorescences with many robbed flowers. The proportion of flowers within an inflorescence visited by P. gigas was inversely proportional to the ratio of robbed flowers.     

Floral colour change in Pedicularis monbeigiana (Orobanchaceae)

We examined the effects of the retention of colour-changed flowers on long- and short-distance attractiveness of bumblebees and the likelihood of successive flower visits by bumblebees in Pedicularis monbeigiana. The lower lip changed colour with age from white to purple. Hand geitonogamous pollination significantly reduced seed production. No pollen limitation occurred in this species. Purple-phase flowers contributed minimally to pollinator attractiveness at long distance. The combination of less reproductive flowers with a lower amount of reward and floral colour change enabled plants to direct pollinators to reproductive, highly rewarding white flowers at close range. A high percentage of purple-phase flowers in an inflorescence was associated with a marked reduction in the frequency of successive flower visits to individual plants. We suggest floral colour change in P. monbeigiana may serve as a mechanism for enhancing inter-individual pollen transfer and reducing intra-individual pollen transfer.     

Foraging behavior of three bee species in a natural mimicry system: female flowers which mimic male flowers in Ecballium elaterium (Cucurbitaceae)

Summary The behavior of Apis mellifera and two species of solitary bees which forage in the flowers of monoecious Ecballium elaterium (L.) A. Rich (Cucurbitaceae) were compared. The female flowers of E. elaterium resemble male flowers visually but are nectarless, and their number is relatively smaller. Apis mellifera was found to discriminate between the two genders and to pay relatively fewer visits to female flowers (mean of 30% relative to male flowers) from the beginning of their activity in the morning. The time spent by honeybees in female flowers is very short compared to that spent in male flowers. It is surmised that the bees remember the differences between the flowers where they foraged on the previous days. In contrast, the two species of solitary bees Lasioglossum politum (Morawitz) (Halictidae) and Ceratina mandibularis Fiese (Anthophoridae) visit the female flowers with nearly equal frequencies at the beginning of each foraging day and stay longer in these flowers. Over the day there is a decline in the relative frequency of visits to female flowers and also in the mean time spent in them. The study shows that bees can collect rewards at high efficiency from the flowers of Ecballium elaterium because of their partial discrimination ability and the scarcity of the mimic flowers. It is suggested that the memory pattern of some solitary bees may be different from that of Apis mellifera. It seems that the limited memory and discrimination ability of bees can lead to a high frequency of visits to the mimic flowers during a long flowering season.     

Visits to Manipulated Flowers by Episyrphus balteatus (Diptera: Syrphidae): Partitioning the Signals of Petals and Anthers

In a number of previous studies attention has been directed to the selection on corolla dimensions by pollinator preference, but anthers may also be a signal. This experiment examined the relative importance of petals and anthers in the attraction of male Episyrphus balteatus (Diptera, Syrphidae) to individual oil-seed rape flowers (Bracus rapae oleifera). Rape flowers have four petals and six anthers. Nine treatments were created, of flowers with four, two, or zero petals and six, three, or zero anthers. Twenty males were tested for preference among the experimental flowers by recording the sequence and duration of their visits (which were all for feeding) to the flowers. The number of first visits and the duration of visits on each flower were significantly related to the number of anthers but not the number of petals. There was no correlation between petal area and pollen load of control flowers. It is suggested that Syrphidae may not always attend to petals, and variation in preference of different pollinator taxa should be taken into account when discussing the evolution of floral characters.     

Breeding systems of the Japanese-endemic carnivorous sundew species Drosera makinoi and Drosera toyoakensis (Droseraceae)

In response to unfavorable environmental conditions, some Drosera species in the northern hemisphere do not open their flowers but form bud-like flowers. The bud-like flowers exhibit pseudo-cleistogamous behavior and produce similar numbers of seeds as open flowers. However, it is not clear how common this type of reproduction is in the genus Drosera. In this study, we examined the effects of flowering (opening flowers/forming bud-like flowers) and pollinator visitation on seed production in Drosera makinoi and D. toyoakensis, two endemic species in Japan. D. makinoi and D. toyoakensis plants did not open 20%–40% chasmogamous flowers to form bud-like flowers. The numbers of mature seeds produced by bud-like flowers were significantly lower than those produced by open flowers (0%–2% of open flowers). In D. makinoi, the percentage of bud-like flowers correlated negatively with the seed production per plant, suggesting that the production of bud-like flowers decreased the lifetime reproductive success. In both D. makinoi and D. toyoakensis, pollinator-excluded flowers produced similar numbers of seeds as control flowers. Our results suggest that pseudo-cleistogamy does not occur in D. makinoi and D. toyoakensis, and they produce seeds without pollinator visits once flowers open.     

Foraging and pollination behaviour of Xylocopa calens Lepeletier (Hymenoptera: Apidae) on Phaseolus coccineus L. (Fabaceae) flowers at Yaounde (Cameroon)

To evaluate impact of carpenter bee, Xylocopa calens, on pod and seed set of Phaseolus coccineus, its foraging and pollinating activities were studied in Yaounde, for two seasons (May–July 2008 and April–June 2009). Observations were made on 40 inflorescences per treatment. The treatments included unlimited floral access by all visitors, bagged flowers to deny all visits, and limited visits by X. calens only. In addition, all flower visitors were recorded. The carpenter bee's seasonal rhythm of activity, its foraging behavior on flowers, its pollination efficiency, the fructification rate and the number of seeds per pod were recorded. Individuals from 16 species of insects were recorded visiting flowers of P. coccineus in the 2 years. Xylocopa calens was the most frequent, followed by Chalicodoma cincta cincta and Apis mellifera. Apart from bees, wasps were also recorded as likely predators. Xylocopa calens mainly foraged for nectar resources. The mean foraging speed was 9.62 flowers/min. Flowers visited by X. calens had higher fruiting rate compared with others, while those bagged had the lowest. In addition, seed formation was higher in X. calens‐visited flowers compared with all others. The results show that this crop experiences pollination deficit even under normal circumstances, considering that flowers visited by X. calens had higher yields compared with those under unlimited access by all visitors. The fruiting rate, the number of seeds/pod and the percentage of normal seeds of unprotected inflorescences were significantly higher than those of inflorescences protected from insects. X. calens foraging resulted in a significant increment of the fruiting rate by 25.80%, as well as the number of seeds/pod by 14.97% and the percentage of normal seeds by 27.75% in 2008 and 18.39% in 2009. Conservation of X. calens nests close to P. coccineus fields could be recommended to improve pod and seed production in the region.     

Solitary and social bees as pollinators of Wahlenbergia (Campanulaceae): single-visit effectiveness, overnight sheltering and responses to flower colour

Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

The pollination ofStenorrhynchos lanceolatus (Aublet) L. C. Rich. (Orchidaceae: Spiranthinae) by hummingbirds in southeastern Brazil

Hummingbird pollination is documented for a natural population ofStenorrhynchos lanceolatus Aublet. L. C. Rich. occurring in Rio de Janeiro State, southeastern Brazil. At the study site the plants are pollinated byPhaethornis eurynome (Phaethorninae),Thalurania glaucopis (females only) andLeucochloris albicollis (Trochilinae). The plants offer nectar as a reward and the pollinaria become stuck to the surface of the hummingbird's bill while it is probing the flowers. The orchid population received a few (0–4) hummingbird visits per day, with about 83% of the flowers being pollinated. In spite of the low frequency of visits, the granular structure of the pollinarium plus the behaviour of the most frequent pollinators, which tend to visit all the fresh-looking flowers of each inflorescence, a very high fruiting success was promoted. Experimental evidence suggests that the pollinaria may remain up to 6.30 hours on the hummingbird's bill, enhancing the chances of cross-pollination and long-distance pollen flow.     

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.     

Florivory and floral larceny by fly larvae decrease nectar availability and hummingbird foraging visits at Heliconia (Heliconiaceae) flowers

Insect larvae inhabit the corolla tubes of some Heliconia species (Heliconiaceae). In this study, we present the first evidence of the influence of these larvae on the pollination ecology of Heliconia plants. We provide experimental evidence that the flowers of Heliconia spathocircinata infested by flies have less nectar for pollinators and received fewer visits by hummingbird pollinators, in comparison with uninfested flowers.     

AUTOGAMY AND COMPETITION FOR POLLINATORS IN HEPATICA AMERICANA (RANUNCULACEAE)

Hepatica americana is one of the earliest flowering vernal herbs in the deciduous forests of piedmont North Carolina. Unlike most other members of the spring wildflower community, its flowers are nectarless and autogamous. The number of insect visits a flower receives varies markedly through the blooming season and is lowest when H. americana is in full bloom. Potential for outcrossing as measured by the seed set of emasculated flowers follows the same pattern. The mid-season decline in insect-pollination is most likely the result of competition with Erythronium umbilicatum, which flowers concurrently with H. americana and is much more attractive to the solitary bees that are the principal early season pollen vectors. Because of this competitive disadvantage, seed set of H. americana in piedmont North Carolina would be pollination-limited if the flowers were not autogamous. Autogamy may also be favored by flowering early in the spring before pollinators are available. Opportunities for outcrossing in H. americana are primarily the result of protogyny and blooming earlier than competitors.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

FLOWER VISITATION,POLLEN DEPOSITION,AND POLLEN-TUBE COMPETITION IN HIBISCUS MOSCHEUTOS (MALVACEAE)

The potential influence of pollen-tube competition on offspring “quality” has received considerable attention in recent years. Yet the prevalence of pollen competition in natural populations is largely unknown because few investigators have actually measured rates of pollen deposition on stigmas. In this study, we assess the potential for pollen-tube competition in natural populations of the self-compatible, pollinator-dependent herbaceous perennial, Hibiscus moscheutos. Individual flowers averaged two to four visits per 15 min by potential pollinators (Ptilothrix and Bombus), and about 34% of these visits involved contact with a stigma. The median number of pollen grains deposited on virgin stigmas per contact visit was 70 grains (values ranged from 0 to 889), and flowers averaged about four contact visits per hour. Approximately 360 pollen grains must reach stigmas for full seed set to occur in a typical flower (an average ovary has 139 ovules, and 2.6 pollen grains are required per seed). Within 2 and 3 hr exposure to pollinators, 65% and 97% of the flowers received excess pollen (>360 grains) and median stigmatic pollen loads exceeded the number of ovules by a factor of 4.0 and 5.4, respectively. Based on 3 yr observations, it is concluded that pollen competition may frequently occur in this species.     

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.     

ANATOMY AND DEVELOPMENT IN SOLISIA

Boke , Norman H. (U. Oklahoma, Norman.) Anatomy and development in Solisia. Amer. Jour. Bot. 47(1): 59—65. Illus. 1960.–The genus Solisia contains a single species of small cacti which resemble certain mammillarias in having: pink, lateral flowers; milky juice; and dimorphic areoles. Adult specimens have elongate spiniferous areoles with an unusual sequence of spine initiation. The first four or five primordia appear at the posterior end of the areole meristem; the next are initiated near its center, after which initiation proceeds both acropetally and basipetally until the single, elliptical series of primordia is complete. A similar pattern of spine initiation occurs in Pelecyphora aselliformis, but this species differs markedly from S. pectinata in other respects. In seedlings of S. pectinata the areoles are broadly elliptical and spine initiation is strictly acropetal, a situation found in certain species of Mammillaria. Seeds of S. pectinata are black with a large hilum and a small perisperm. Since the perisperm has apparently been overlooked, it appears that only the black seed coat and relatively large hilum keep the species out of Mammillaria. If Buxbaum's postulates concerning the value of seed structure in tracing phylogeny in the Cactaceae are valid, S. pectinata must have diverged from the main line of evolution somewhere below Neobesseya. In that event, the species probably merits generic rank; otherwise, it seems preferable to return it to Mammillaria.     

Nectar robbing of a carpenter bee and its effects on the reproductive fitness of Glechoma longituba (Lamiaceae)

The floral biology and pollination process of Glechoma longituba (Nakai) Kuprian, a clonal gynodioecious, perennial herb that is widely distributed in China was investigated in natural populations. During visits to the flowers of G. longituba, the carpenter bee—Xylocopa sinensis mainly displayed nectar-robbing behavior with minimal pollination. Nectar robbing behavior began at the onset of flowering and continued for about 3 weeks ending at about the middle of the flowering period. A total of 18.6% floral visits in this period were by nectar robbers, with about 90% of the flowers in the study populations being subjected to two or two nectar-robbing episodes. During controlled experiments, lower pollination efficiency was recorded for X. sinensis compared to legitimate pollinators. Pollination by the robber-like pollinator X. sinensis during the early-mid phase of the flowering season yielded seeds of 16.16%. Although nectar robbing by the carpenter bee seemed to have a slight enhancing effect on seed set in G. longituba, this effect was effectively masked by the more pronounced detrimental effect of nectar robbing. Experiments indicated that nectar robbing by the carpenter bee reduced seed production by more than 26%, largely owing to the consequent shortening of the life span of robbed flowers. Furthermore, 10.43% of the ovules and 13.04% of the nectaries in the robbed flowers were damaged, thus causing a decrease or entire loss of reproductive opportunity in the robbed flowers. In addition, a higher number of pollen grains remained on the androecia of robbed flowers indicating that nectar robbing may have a lowering effect on male fitness in G. longituba.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

Primary nectar robbing by Apis mellifera (Apidae) on Pyrostegia venusta (Bignoniaceae): behavior,pillaging rate,and its consequences

The interactions between plants and their pollinators are the result of convergent evolution of floral attributes reflecting pressure exerted by pollinators. Nonetheless, the strategies employed by floral visitors to collect floral resources are extremely complex, and commonly involve theft or robbery in addition to pollination. We describe here the behavioral repertory of Apis mellifera during the collection of the floral resources, and evaluated the robbing rates of A. mellifera on the buds and flowers of Pyrostegia venusta during periods of intense and sparse flowering. We recorded the behaviors exhibited by foraging bees while collecting floral resources, quantified the numbers of floral buds and flowers with perforations in their corolla tissues, and determined whether that damage reduced nectar production. The evaluations were conducted during two distinct periods: during the period of intense flowering of P. venusta, and during the period of sparse flowering. Nectar robbing was observed during 93.4% of the visits of foraging A. mellifera bees, while nectar theft was observed during only 0.7% of the visits, and pollen theft during 5.9%. The robbing of floral buds and flowers was most intense during the period of heavy flowering. Flowers that had been intensely robbed secreted significantly less nectar than those non-robbed. The unusual nectar robbing activities of A. mellifera, especially during the period of intense flowering indicates an optimization of access to larger volumes of food resources. Our results therefore point to a major limitation of nectar per floral unit during the intense flowering period of P. venusta due to the high activity of nectar robbing by A. mellifera bees.

     

FLORAL MORPHOLOGY AND CROSS-POLLINATION IN ERYTHRONIUM GRANDIFLORUM (LILIACEAE)

In bumblebee visits to flowers of Erythronium grandiflorum (Liliaceae), the ratio of self- to nonself- (“outcross”) pollen grains deposited on the stigma is positively correlated with the degree of stylar exsertion beyond the anthers. Natural populations show substantial, continuous variation in stylar exsertion.