Comparative nectar-foraging behaviors and efficiencies of an alien and a native bumble bee

Resource preemption by alien organisms can contribute to their invasion success and the demise of functionally equivalent native species, particularly when opportunistic foraging by aliens results in more efficient exploitation. In forests of NW Patagonia, the only native bumble bee and major pollinator, Bombus dahlbomii, declined almost to extinction as the alien B. ruderatus increased in abundance since its first appearance about 17 years ago. To explore whether resource competition might have driven this displacement we studied the behavior and foraging efficiency of both bumble bees while they harvested nectar from flowers of Alstroemeria aurea, the main summer food resource in the forests of NW Patagonia. We compared the nectar content of flowers that bees selected, recently visited, and rejected with that of randomly-chosen neighboring flowers and assessed differences in visitation rates. The native bumble bee selects flowers with abundant nectar and mostly exploits nectar-rich flower patches by rejecting a higher proportion of flowers with little or no nectar. On the other hand, the alien bumble bee discriminated less with respect to sugar content per visited flower, but visited more flowers per minute. Workers of the native bumble bee harvested ~70% more sugar per unit of time than those of the alien species in absolute terms, and a similar amount when sugar harvested was expressed as a percentage of body mass. In contrast to expectation, the opportunistic foraging of the alien bumblebee was not more efficient and therefore cannot explain the ecological extinction of the native species through exploitative competition. These findings suggest that the displacement of the native species by the alien may be driven by other factors, such as the associated introduction of novel diseases or parasites.     

Community-dependent foraging habits of flower visitors: cascading indirect interactions among five bumble bee species

Despite the ubiquity and the importance of interspecific interactions among flower visitors, few studies have examined their effects on the realized feeding niches of visitor species in a community context. To evaluate the community-wide effects of interactions among flower visitors, I have examined changes in the flower utilization patterns of each visitor species at several sites where the component of the visitor’s community differed. Specifically, I compared the flower preferences and foraging habits (legitimate foraging vs. primary nectar robbing vs. secondary nectar robbing) of five bumble bee species in flower patches consisting of Trifolium pratense L. (red clover) and T. repens L. (white clover) on Hokkaido Island, Japan. I also examined the nectar production and standing crops of each flower species to evaluate the exploitation competition based on nectar. The bumble bee species exhibited different flower utilization patterns among sites. At sites where the long-tongued Bombus diversus tersatus was common and the exotic short-tongued B. terrestris was rare, B. diversus tersatus visited red clover (long-tubed flowers) exclusively, whereas medium-tongued B. pseudobaicalensis and short-tongued B. hypocrita sapporoensis and B. hypnorum koropokkrus preferentially visited white clover (short-tubed flowers). Conversely, at sites where the long-tongued bee was rare, four other species frequently visited red clover in different modes: B. pseudobaicalensis visited legitimately, B. hypocrita sapporoensis and B. terrestris visited as primary nectar robbers, and B. hypnorum koropokkrus visited as a secondary nectar robber. The presence or absence of resource exploitation by the long-tongued species and the interaction between primary and secondary nectar robbers via robbing holes was the major ecological sources of these differences. Diverse effects of interactions among flower visitors played important roles in shaping pattern of plant and flower visitor interactions.     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Bumblebees Do Not Respond to Variance in Nectar Concentration

Worker bumblebees (Bombus fervidus) were given repeated binary choices between two colors of artificial flowers with the same associated mean nectar concentration (X? = 20%), but with different variances in nectar concentration. Flowers of one color, yellow or blue, rewarded a bee with 1 μl of 20% sucrose solution (low-variance flower type) on each visit (p = 1) and flowers of the other color rewarded a bee on each visit with 1 μl of either 10% or 30% sucrose (p = 0.5; high-variance flower type). Of the 10 bees tested, nine showed no preference for either the high- or low-variance flowers (indifferent or risk-insensitive). This result is similar to honeybee responses to variation in nectar concentration, despite differences in foraging ecology between bumblebees and honeybees. Flower-choice behaviour in the presence of variance in nectar concentration is a response to the expected concentration of the alternative flower types. Possible mechanisms of risk-sensitive foraging behaviour in bees are discussed.     

Aggressive foraging of social bees as a mechanism of floral resource partitioning in an Asian tropical rainforest

Interference competition by aggressive foraging often explains resource partitioning, but mechanisms contributing to partitioning have rarely been studied in Asian social bee guilds. Foraging of social bees at canopy flowers of Santiria laevigata (Burseraceae) and honey-water feeders was studied in a lowland mixed-dipterocarp forest in Sarawak, Malaysia. Four stingless bee species (Apidae, Meliponinae), Trigona canifrons, T.␣fimbriata, T. apicalis and T. melina, aggressively defended flower patches and feeders. At the flowers, T.␣canifrons excluded other bees only in the morning when nectar flow peaked. At the feeders, the aggression resulted in asymmetric interference competition, which produced a dominance hierarchy among seven social bee species. Interspecific partitioning of the feeders was detected in time and height but not quality. Only time of the first arrival after feeder presentation was negatively correlated with the dominance hierarchy: more aggressive species arrived at the feeders later than less aggressive species. This result suggests that a trade-off between searching ability and defensive ability at flower patches gives rise to resource partitioning in the social bee guild. Received: 22 June 1996 / Accepted: 22 November 1996     

A Field Study of Spatial Memory in Green-Backed Firecrown Hummingbirds (Sephanoides sephaniodes)

The foraging ecology of hummingbirds involves the exploitation of a high number of patchily distributed flowers. This scenario seems to have influenced capabilities related to learning and memory, which help to avoid recently visited flowers and to allocate exploitation to the most rewarding flowers, once learning has occurred. We carried out two field experiments with the green‐backed firecrown hummingbird (Sephanoides sephaniodes, Trochilidae) in order to examine the ability of birds, first, to recall a nectar location, and secondly, to remember the location of the most rewarding flower among lower quality flowers. The first experiment showed that subjects were able to recall the location of nectar among flowers of identical appearance. In the second experiment, hummingbirds were also able to recall the location of the most rewarding nectar among less rewarding flowers with the same appearance. The results of this study suggest that S. sephaniodes can remember the location of the most rewarding patch, facilitating efficient exploitation of flowers in the absence of visual cues related to nectar quality.     

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.     

Temporal and microclimatic partitioning of the floral resources of Justicia aurea amongst a concourse of pollen vectors and nectar robbers

Summary The flowers of Justicia aurea, morphologically characteristic of ornithophily, attracted a diverse array of foragers where they occurred as a dense stand in the tropical forests at La Selva, Costa Rica, and so provided an arena for this study of competition and coexistence. Two hummingbird species (Phaethornis superciliosus and Campylopterus hemileucurus) visited the flowers legally early in the morning, and defended the nectar resource; a third smaller bird (P. longuemareus) foraged for nectar throughout the day but collected it illegally by piercing the corollas. In addition, nectar was harvested illegally by four species of stingless bee (Trigona) and by ants, creating a further drain on the limited floral resources.Consideration of the diurnal patterns of foraging activities in combination with a spatial axis (defined here in terms of microclimate and insolation) nevertheless showed a good separation of flower visits for the different nectarivores. Hummingbirds visited flowers in zones where the reward was highest, while insects foraged to minimise their energetic costs; each of these factors could in turn be related to microclimatic considerations. A comprehensive scheme of resource utilisation could therefore be extracted from the field observations and interpreted in these terms. The limited area of niche overlap thus revealed corresponded closely with the situations where direct interference competition was observed, between hummingbird species or between bees and aggressive ants.     

Morphological variation in relation to flower use in bumblebees

To understand resource partitioning in a bumblebee community, we analyzed various morphological characters. A total of 1269 individuals of six bumblebee species, Bombus ardens, B. hypocrita, B. diversus, B. ignitus, B. honshuensis and B. beaticola, were examined and principal component analysis showed that the bumblebee species were clearly differentiated. Glossa, prementa and head lengths were positively correlated with the second component, and a longer proboscis was associated with a narrower body, which may help bees to intrude into and access deep‐lying nectar sources. Bombus diversus, with a long proboscis and narrow body, preferred flowers with a long corolla tube, whereas B. hypocrita and B. ignitus, which have short proboscises and wide bodies, visited flowers with short corollas or dish‐shaped flowers. Two pairs of consubgeneric species that have similar morphological characteristics, B. ardens and B. beaticola, and B. hypocrita and B. ignitus, divided flower resources by habitat selection and seasonal partitioning. For resource partitioning among bumblebee species, not only morphology but also other factors, such as habitat and seasonal preference, flower use, foraging behavior, and interspecific interactions, are responsible.     

Monitoring Flower Visitation Networks and Interactions between Pairs of Bumble Bees in a Large Outdoor Flight Cage

Pollinators, such as bees, often develop multi-location routes (traplines) to exploit subsets of flower patches within larger plant populations. How individuals establish such foraging areas in the presence of other foragers is poorly explored. Here we investigated the foraging patterns of pairs of bumble bees (Bombus terrestris) released sequentially into an 880m² outdoor flight cage containing 10 feeding stations (artificial flowers). Using motion-sensitive video cameras mounted on flowers, we mapped the flower visitation networks of both foragers, quantified their interactions and compared their foraging success over an entire day. Overall, bees that were released first (residents) travelled 37% faster and collected 77% more nectar, thereby reaching a net energy intake rate 64% higher than bees released second (newcomers). However, this prior-experience advantage decreased as newcomers became familiar with the spatial configuration of the flower array. When both bees visited the same flower simultaneously, the most frequent outcome was for the resident to evict the newcomer. On the rare occasions when newcomers evicted residents, the two bees increased their frequency of return visits to that flower. These competitive interactions led to a significant (if only partial) spatial overlap between the foraging patterns of pairs of bees. While newcomers may initially use social cues (such as olfactory footprints) to exploit flowers used by residents, either because such cues indicate higher rewards and/or safety from predation, residents may attempt to preserve their monopoly over familiar resources through exploitation and interference. We discuss how these interactions may favour spatial partitioning, thereby maximising the foraging efficiency of individuals and colonies.     

Risk‐averse inflorescence departure in hummingbirds and bumble bees: could plants benefit from variable nectar volumes?

Most hermaphroditic, many-flowered plants should suffer reduced fitness from within-plant selfing (geitonogamy). Large inflorescences are most attractive to pollinators, but also promote many flower visits during a single plant visit, which may increase selfing and decrease pollen export. A plant might avoid the negative consequences of attractiveness through modification of the floral display to promote fewer flower visits, while retaining attractiveness. This report shows that increasing only the variance in nectar volume per flower results in fewer flower visits per inflorescence by wild hummingbirds ( Selasphorus rufus ) and captive bumble bees ( Bombus flavifrons ) foraging on artificial inflorescences. Inflorescences were either constant (all flowers contained the same nectar volume) or variable (half the flowers were empty, the other half contained twice as much nectar as in the constant flowers). Both types of inflorescence were simultaneously available to foragers. Risk-averse foraging behaviour was expressed as a patch departure preference: birds and bees visited fewer flowers on variable inflorescences, and this preference was expressed when resource variability could be determined only by concurrent sampling. When variance treatments were clearly labelled with colour and offered to hummingbirds, the departure effect was maintained; however, when preference was measured by inflorescence choice, birds did not consistently prefer to visit constant inflorescences. The reduced visitation lengths on variable inflorescences by both birds and bees documented in this study imply that variance in nectar production rates within inflorescences may represent an adaptive trait to avoid the costs of geitonogamy.     

Specialised protagonists in a plant–pollinator interaction: the pollination of Blumenbachia insignis (Loasaceae)

• Analyses of resource presentation, floral morphology and pollinator behaviour are essential for understanding specialised plant‐pollinator systems. We investigated whether foraging by individual bee pollinators fits the floral morphology and functioning of Blumenbachia insignis, whose flowers are characterised by a nectar scale‐staminode complex and pollen release by thigmonastic stamen movements.
• We described pollen and nectar presentation, analysed the breeding system and the foraging strategy of bee pollinators. We determined the nectar production pattern and documented variations in the longevity of floral phases and stigmatic pollen loads of pollinator‐visited and unvisited flowers.
• Bicolletes indigoticus (Colletidae) was the sole pollinator with females revisiting flowers in staminate and pistillate phases at short intervals, guaranteeing cross‐pollen flow. Nectar stored in the nectar scale‐staminode complex had a high sugar concentration and was produced continuously in minute amounts (~0.09 μl·h^?1). Pushing the scales outward, bees took up nectar, triggering stamen movements and accelerating pollen presentation. Experimental simulation of this nectar uptake increased the number of moved stamens per hour by a factor of four. Flowers visited by pollinators received six‐fold more pollen on the stigma than unvisited flowers, had shortened staminate and pistillate phases and increased fruit and seed set.
• Flower handling and foraging by Bicolletes indigoticus were consonant with the complex flower morphology and functioning of Blumenbachia insignis. Continuous nectar production in minute quantities but at high sugar concentration influences the pollen foraging of the bees. Partitioning of resources lead to absolute flower fidelity and stereotyped foraging behaviour by the sole effective oligolectic bee pollinator.

     

Observational Learning in the White-Eared Hummingbird (Hylocharis leucotis): Experimental Evidence

The adoption of new food resources can be facilitated by the ability to learn through observation of other individuals who use them. This behavior, termed observational learning, applies to any problem solving in which a naive individual who has observed an experienced individual learns a behavior faster than another who has not. Hummingbirds consume nectar from flowers of a large number of plant species, which are very diverse in morphology and color. During their local or migratory movements, they can observe the use of floral resources by conspecifics and heterospecifics which may change their foraging preferences. Although there is evidence that hummingbirds can use observational learning to exploit new floral resources, it is necessary to generate additional information by studying different hummingbird species. In this work, the learning performance of White‐eared hummingbirds (Hylocharis leucotis) was studied in the presence or absence of a knowledgeable tutor. In a first experiment, naïve hummingbirds learned to feed on arrays of artificial flower of two colors: red (previously known resource) and yellow (novel resource), where only one color had nectar. Naive hummingbirds visited red flowers faster and more often than rewarded yellow flowers. Individuals with the best performance on each color were further trained to ensure that they only visited flowers of a specific color, and were then used as tutors in the next experiment, in which new naive hummingbirds, caged individually, were allowed to observe them foraging on the artificial arrays. These naïve individual were then exposed alone to the same array used by their tutor. Tutored hummingbirds learned to feed faster and more frequently from nectar‐containing flowers of the array than naive individuals. Likewise, all tutored individuals only visited flowers of the color that had been previously visited by their tutors. This study provides experimental evidence that hummingbirds taken directly from the field can use observational learning as an efficient strategy to access new floral resources.     

西方蜜蜂和兰州熊蜂在设施桃园的访花偏好性比较

【目的】为了比较西方蜜蜂 Apis mellifera 和兰州熊蜂 Bombus lantschouensis 在设施桃园内对不同时期桃花的访花偏好性、以及这种偏好性与花粉活力和采集花粉花蜜之间的关系。【方法】记录2种蜂在温室桃园内访问早期花、中期花和晚期花的比例,测定桃花不同时期的花粉活力以及2种蜂携带的花粉活力,观察2种蜂采集花蜜和采集花粉的成功率,统计2种蜂访花过程中桃花所处的枝条数及植株数。【结果】桃花不同时期的花粉活力差异显著,早期花花药未开裂,花粉未释放,中期花花粉活力（58.3%）显著高于晚期花花粉活力（34.2%）（P<0.01）;西方蜜蜂更加偏好访问中期花,对中期花的访问率高达75.3%,显著高于兰州熊蜂对中期花的访问率（49.2%）（P<0.01）;西方蜜蜂携带的花粉活力（92.1%）显著高于兰州熊蜂携带的花粉活力（72.9%）,但是西方蜜蜂采集花粉和采集花蜜的成功率均低于兰州熊蜂（P<0.01）;在访问一定数量的桃花过程中,兰州熊蜂在设施桃园内访问的枝条数和植株数较多,分布范围较广（P<0.01）。【结论】和兰州熊蜂相比,西方蜜蜂对活力花粉的辨别能力更强,更加偏好访问花粉活力较高的花朵,这种偏好性导致其采集花粉花蜜的成功率降低。     

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.     

Nectar secretion dynamic links pollinator behavior to consequences for plant reproductive success in the ornithophilous mistletoe Psittacanthus robustus

The mistletoe Psittacanthus robustus was studied as a model to link flower phenology and nectar secretion strategy to pollinator behaviour and the reproductive consequences for the plant. The bright‐coloured flowers presented diurnal anthesis, opened asynchronously throughout the rainy season and produced copious dilute nectar as the main reward for pollinators. Most nectar was secreted just after flower opening, with little sugar replenishment after experimental removals. During the second day of anthesis in bagged flowers, the flowers quickly reabsorbed the offered nectar. Low values of nectar standing crop recorded in open flowers can be linked with high visitation rates by bird pollinators. Eight hummingbirds and two passerines were observed as potential pollinators. The most frequent flower visitors were the hummingbirds Eupetomena macroura and Colibri serrirostris, which actively defended flowering mistletoes. The spatial separation between anthers, stigma and nectar chamber promotes pollen deposition on flapping wings of hovering hummingbirds that usually probe many flowers per visit. Seed set did not differ between hand‐, self‐ and cross‐pollinated flowers, but these treatments set significantly more seeds than flowers naturally exposed to flower visitors. We suggest that the limitation observed in the reproductive success of this plant is not related to pollinator scarcity, but probably to the extreme frequency of visitation by territorial hummingbirds. We conclude that the costs and benefits of plant reproduction depend on the interaction strength between flowers and pollinators, and the assessment of nectar secretion dynamics, pollinator behaviour and plant breeding system allows clarification of the complexity of such associations.     

The foraging behaviour of Australian honeyeaters: a review and some comparisons with hummingbirds

The foraging behaviour of Australian honeyeaters is reviewed in terms of diet, foraging selectivity, foraging flight mode, quality and quantity of nectar encountered per flower, flower densities encountered and effect of predation. At the same time comparisons are made between honeyeaters and hummingbirds. These two groups of birds are superficially similar. Both feed on nectar and insects. Both tend to have long curved bills and tongues adapted for removal of nectar from flowers. Both tend to feed at long, red flowers. However, on close inspection, honeyeaters and hummingbirds are quite dissimilar. For example, many honeyeaters include fruit in their diets. Hummingbirds almost never eat fruit. Honeyeaters appear to be considerably less nectarivorous and more insectivorous than hummingbirds. Honeyeaters are, for the most part, larger than hummingbirds and they usually perch while feeding whereas hummingbirds usually hover. Honeyeaters but not hummingbirds often flock while feeding. Predation appears to be considerably more important for honeyeaters than for hummingbirds. Territorial defense of flowers seems common in hummingbirds but uncommon in honeyeaters. These differences are discussed in detail and explanations are offered for them wherever possible.     

The pollination of Bromelia antiacantha (Bromeliaceae) in southeastern Brazil: ornithophilous versus melittophilous features

Bromelia antiacantha flowered from December to February and during this period the central leaves and bracts displayed a bright red colour. The inflorescence bears 150-350 flowers, with 10-35 flowers opening per day over 4-5 days. The flowers are dark magenta coloured with white margins, tubular-shaped with a wide opening, and their stigma is situated below the anthers. Anthesis began around 4:00 h and flowers lasted approximately 15 h. The highest nectar volume and sugar concentration occurred between 4:00-6:00 h; after this period, both decreased throughout the day. B. antiacantha is partially self-incompatible, non-autogamous, and therefore, pollinator dependent. The hummingbirds Thalurania glaucopis, Amazilia fimbriata, and Ramphodon naevius were its most frequent pollinators (55% of the visits), visiting flowers mainly in the afternoon. The scattered distribution of B. antiacantha promoted trap-lining behaviour of the hummingbirds, which favoured fruit set through xenogamy. Corolla colour, wide flower opening, sweet odour and concentrated nectar early in the day favoured bee visitation. Of the 38 % of bee visits, 96% were made by Bombus morio, mainly in the morning and their behaviour promoted self-pollination. The bee Trigona spinipes frequently acted as a pollen and nectar thief early in the morning, excluding most of the hummingbirds. Bromelia antiacantha has ornithophilous and melittophilous features, and despite being pollinated by two kinds of agents, its reproductive success depends on a given set of circumstances.     

Mobility of Impatiens capensis flowers: effect on pollen deposition and hummingbird foraging

Flexible pedicels are characteristic of birdpollinated plants, yet have received little attention in studies of hummingbird-flower interactions. A major implication of flexible pedicels is that flowers may move during pollination. We examined whether such motion affected interactions between ruby-throated hummingbirds (Archilochus colubris) and jewelweed (Impatiens capensis) by increasing pollen deposition and by altering the effectiveness of nectar removal. For I. capensis, flower mobility enhanced pollen deposition: birds had significantly longer contact with anthers and more pollen deposited on their bills and crowns when foraging at mobile flowers than at flowers that had been experimentally immobilized. In contrast, flower mobility imposed a cost on hummingbirds by significantly increasing their handling times and reducing their extraction rates relative to their interactions with immobile flowers. Field observations indicated that the motion observed during hummingbird visits did not occur when bees (Bombus spp., Apis mellifera) visited I. capensis flowers, which suggests that the mobility of I. capensis flowers is an adaptation for hummingbird pollination.     

Pollination ecology of Penstemon roseus (Plantaginaceae), an endemic perennial shifted toward hummingbird specialization?

Summary Pink-flowered tubular Penstemon roseus (Plantaginaceae), which has shifted partially to hummingbird pollination, blooms on high-elevation slopes in the mountains in Tlaxcala, Mexico. We studied the interactions between pollinator visitation rates to flowers, pollen removal and deposition, flower size, and nectar removal frequency on seed production in P. roseus. We combine observational and experimental studies in two contrasting natural populations. Our manual pollinations revealed that P. roseus is fully self-compatible. Autonomous self- and manual self-pollinated flowers matured as many seeds as when outcrossed, but outcrossing seems to become better than selfing as the flowering season progressed. Early in the season flowers that were bagged and hand-selfed, hand-outcrossed, or autonomously selfed, or unbagged and naturally pollinated had equal seed set in all four treatments. But later in the season, outcross pollen gave approximately twice as much seed set as the two self-treatments. Low levels of pollen receipt and pollen removal were consistent with the long time elapsed for a given plant to be visited by hummingbirds, which suggests pollen shortage in both sites. Despite differences in pollinator visitation rates to flowers, probability of flower visitation, removal and deposition of pollen, and nectar production rates between populations, we found that total nectar production had no effect on seed production at either site. The daily nectar secretion rate of 0.3–0.65 mg sugar per flower per 1–3 days was low relative to other hummingbird-adapted Penstemon species (typical range: 1.5–5 mg sugar per flower), and it might be intermediate between hummingbird- and bee-adapted Penstemon flowers. Our results support the hypothesis about a shift toward hummingbird pollination, and provide an example of a ‘despecialized’ Penstemon species, which attracts high-energy pollinators (hummingbirds) and profits from outcrossing, but retains bee-syndrome floral traits and low sugar production rates.