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1.
Nectar-feeding birds are prominent in many parts of the world, and vary with respect to body size. Despite the availability of considerable morphometric data, few concerted efforts have been made to assess the influence of attributes such as mass, wing length and leg morphology upon the speed, acceleration, mode and energetic cost of movement by birds between flowers when foraging for nectar. This review attempts to consolidate and interpret available data and highlight areas where further investigations appear warranted. Australian honeyeaters are generally larger, and American hummingbirds smaller, than Hawaiian honeycreepers and sunbirds of Africa or Asia. Sunbirds, honeyeaters and honeycreepers generally perch while extracting nectar from flowers. Hummingbirds usually hover, apparently because suitable perches close to flowers are lacking, and not because hovering increases the speed at which flowers can be visited. Honeyeaters move from one flower to another at speeds that are at least as great as those for hummingbirds. Most passerine nectarivores need to ingest more nectar per day than hummingbirds in order to maintain energy balance, some species devoting more than 60% of the day to foraging. The major consequence of reduced foraging activity by hummingbirds, which spend only 5–30% of the day in this manner, appears to be male emancipation from nest construction and care of offspring. Large nectarivores have a greater capacity to store surplus food and to fast than smaller birds, and so can take advantage of short-lived peaks in nectar abundance. Nectarivores such as honeyeaters should therefore be favoured by the rapid diurnal changes in nectar availability which are characteristic of many Australian and African habitats. Body mass also determines the likely access to rich sources of nectar through size-related interspecific dominance hierarchies. In all families, larger species tend to monopolize the most rewarding nectar supplies, forcing smaller subordinate species to use poorer, more scattered sources. Within particular species, males usually have longer wings and greater masses than females. These variations imply that the two sexes differ with regard to their foraging ecology, although few supporting data are currently available.  相似文献   

2.
Nectar availability in Calothamnus quadrifidus flowers was studied at Wongamine Nature Reserve in late spring (November). Despite some overnight depletion by moths and other invertebrates, more nectar was present in flowers at dawn than at the preceding dusk. Significant nectar depletion occurred within a few hours after dawn, mainly due to foraging by two honeyeater species. Lichmera indistincta and Phylidonyris nigra. Thereafter, nectar availability was maintained at relatively low levels, principally because of foraging by honeyeaters and honey bees. Apis mellifera, that became active during the warmer part of the day. Although individual honeyeaters consumed more nectar than A. mellifera, honey bees were so abundant that their total impact was greater than that of either honeyeater species for much of the day. Transfer of C. quadrifidus pollen between flowers is necessary in order to achieve a high level of seed set, as the flowers appear to be protandrous. Honeyeaters appeared to be considerably more significant pollen vectors than A. mellifera.  相似文献   

3.
The bills and tongues of nectar-feeding birds differ from continent to continent. The major differences are that: (i) the tongues of A Australian honeyeaters are broader any more fimbricated at the tip than the bifurcated tongues of sunbirds and hummingbirds; (ii) the bills of hummingbirds are more uniformly narrow and taper less markedly towards their tips than those of sun-birds and honeyeaters; and (iii) bill curvatures are generally greater for sunbirds and honey-creepers than for hummingbirds. A variety of hummingbirds has straight or even slightly upturned bills, while bills for all sunbirds, honeycreepers and honeyeaters are decurved to some extent. Despite differences in tongue morphology, hummingbirds, sunbirds and honeyeaters extract nectar at a similar range of rates, averaging approximately 40 γL s?1 from ad libitum feeders, and 1–15 γL?1 from flowers. All tongues collect nectar by capillarity, with licking rates of 6–17 s?1. Licking behaviour has been little studied, although speeds of licking respond to changes in sugar concentration and corolla length. The tongues of honeyeaters are broad, and may need to be brush-tipped in order to allow capillary collection of nectar. Brush-tipped tongues can cover large surface areas on each lick, and may allow honeyeaters to exploit nectar and honeydew that is thinly spread over large surface areas. Bill lengths of nectarivorous birds are similar in all regions, though species of hummingbird have the shortest and longest bills. Bill lengths largely determine the range of floral lengths that can be legitimately probed. Maximum floral lengths exceed bill lengths, since hummingbirds, sunbirds and honeyeaters protrude their tongues beyond the tips of their bills. Rates of nectar extraction, however, decline rapidly once the floral length exceeds bill length. Decurved bills may have evolved in honeyeaters and sunbirds to enable perching birds to reach flowers at the ends of branches more easily. Consistent differences in bill length between the sexes suggest that males and females may exploit different floral resources or different proportions of the same resources. For honeyeaters and sunbirds, males have longer bills than females, but the reverse is true for many hummingbirds.  相似文献   

4.
Honeyeaters (Meliphagidae) were observed foraging for nectar from Lambertia formosa inflorescences, each of which has seven flowers. The frequency distribution of numbers of flowers probed per visit to an inflorescence was found to be bimodal, with one peak at two and the other at seven. It is hypothesized that this frequency distribution results from a rule of departure from inflorescences that maximizes the net rate of energy gain. Patterns of nectar distribution were determined for a large sample of inflorescences. In addition the extent to which the honeyeaters re-probe flowers during a visit to an inflorescence was estimated. From these data and from field measurements of the times required by the honeyeaters to perform the various foraging behaviours, computer simulations of honeyeater foraging were constructed. These simulations led in turn to optimal frequency distributions of numbers of flowers probed per inflorescence that were bimodal but had peaks at 1 and 7 instead of 2 and 7. Although the observed and predicted behaviour were consequently similar, the difference between them was nevertheless significant. This difference could have been due to the birds' transient occupancy of the study area.  相似文献   

5.
Observations of hummingbirds feeding at flowers longer or shorter than their bills seem to contradict the view that bill lengths of hummingbirds evolved in concert with the lengths of their flowers. Recent experiments, however, indicate that a hummingbird's ability to feed at artificial flowers of different lengths depends on the widths of the flowers. We examined if the broad range of flower lengths visited by many hummingbird species can be explained by the widths of the flowers. We predicted that both short‐ and long‐billed hummingbirds would include long, wide flower species in their diets, but that short‐billed hummingbirds would not include long, narrow flower species because nectar in these species might be beyond the reach of their bills. If so, the slope of the regression for flower width versus flower length should be smaller for flower species visited by longer‐billed hummingbirds relative to those visited by shorter‐billed hummingbirds. Analyses of data sets for some North American and Monteverde hummingbirds and their food plants were consistent with this prediction, and bill lengths were significantly correlated with the slopes of the regressions of flower width versus length for seven hummingbird species. Comparisons of observed flower use by some Monteverde hummingbird species to flower assemblages generated at random suggest that these significant regressions were not simply a result of allometric relationships between flower lengths and widths, but in some cases reflected active choice by the birds. The two hummingbird–flower data sets also differed significandy in the scaling of corolla width relative to corolla length. In particular, the Monteverde data set contained a large number of long, narrow flower species, which we suggest is a consequence of a different floral evolutionary history and association with long‐billed hummingbird species. The evolutionary effects of hummingbirds and their flowers upon one another are more complex than has generally been realized, and a consideration of corolla length jointly with other floral characters may improve our understanding of hummingbird‐flower relationships.  相似文献   

6.
Hummingbird flower mites are transported in the nares of hummingbirds and may compete with them by "robbing" nectar secreted by the host plants. We have shown that Tropicoseius sp. flower mites consume almost half the nectar secreted by the long-lived, protandrous flowers of Moussonia deppeana (Gesneriaceae) pollinated by Lampornis amethystinus (Trochilidae). In this paper, we ask whether mimicking nectar consumption of flower mites alters some aspects of hummingbird foraging patterns, and, if so, how this affects host plant seed production. We observed hummingbirds foraging on (a) plants in which nectar was removed from the flowers and then filled with a sugar solution to half the volume of nectar simulating nectar consumption by flower mites, and (b) plants where nectar was removed and then filled with the sugar solution up to normal nectar volumes. Flower mites were excluded from both groups of plants to control for mite activity. Hummingbirds made fewer but longer visits to plants and revisited more the flowers with nectar removal than those without the treatment. We then conducted a pollination experiment on pistillate flowers using a stuffed L. amethystinus hummingbird to evaluate the effect of pollination intensity (number of bill insertions into one flower) on seed production. Flowers with more insertions produced significantly more seeds than those flowers that received fewer insertions. We conclude that the simulation of nectar consumption by hummingbird flower mites can influence the behavior of the pollinator, and this may positively affect seed production.  相似文献   

7.
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.  相似文献   

8.
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.  相似文献   

9.
Abstract

Honeyeaters were the most numerous birds in banksia woodland near Perth, Western Australia, throughout the year. Numbers were greatest in a Banksia littoralis swamp, but only during those few months when it contained large amounts of nectar. In the surrounding woodland, numbers were lower but fairly constant during the year. This reflects the smaller amounts of nectar produced throughout the year, by the overlapping flowering patterns of several Banksia and Adenanthos species.

Large and medium-sized honeyeaters (wattlebirds and New Holland Honeyeaters) and flocking silvereyes dominated the swamp when it flowered. In contrast, small honeyeaters (spinebills and Brown Honeyeaters), many of whom were highly territorial residents, comprised the majority of the woodland assemblage throughout the year. These observations support a model based upon aggressive defence of rich nectar sources by the larger honeyeater species, and more efficient exploitation of dispersed flowers by smaller honeyeaters.  相似文献   

10.
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.  相似文献   

11.
BACKGROUND AND AIMS: Hummingbird flower mites feed and reproduce in flowers of host plants pollinated by hummingbirds, and use the nostrils and bill of the hummingbird to move from plant to plant. These mites compete with the pollinator for the nectar produced by flowers. An investigation was made of the relationship between the pattern of nectar production and the effects of hummingbird flower mites in the flowers of two sympatric species of Heliconia (Heliconiaceae). METHODS: Nectar production was sampled by carrying out two experiments: 2-hour intervals and accumulated nectar. Flowers with and without mites were used in both experiments. KEY RESULTS: Exclusion of mites increased nectar production, especially in accumulated daily production (a maximum of 49 % more nectar). Both Heliconia species had the same pattern of nectar production: a high concentration in the morning, which was progressively reduced as the day passed. This pattern of nectar production coincides with the behaviour of the pollinator, which makes more frequent visits in the morning, as observed in a previous study. CONCLUSIONS: The results suggest that the impact of mites on nectar availability of Heliconia is more important with regard to total volume of nectar produced irrespective of flower longevity. A high variation among individuals in nectar produced in the populations was also observed. Hummingbird flower mites strongly affect availability of nectar for hummingbirds.  相似文献   

12.
Summary During October and November, 1977, a study of nectar production and nectarivore foraging in Eucalyptus incrassata was conducted at Wyperfeld National Park in south-eastern Australia in order to evaluate the extent to which introduced honeybees (Apis mellifera) compete with native honeyeaters for floral nectar. Data on nectar production, nectar availability, ambient air temperature and the numbers of visiting honeyeaters and honeybees were collected. Most of the daily nectar production in E. incrassata occurs early in the morning when temperatures are too low for insects to forage. In addition, insects, particularly honeybees, are unable to exploit nectar in the youngest flowers because the stamens are clustered tightly around the style. As a result of these temporal and structural characteristics of the flowers, honeyeaters are able to harvest most of the nectar. Honeybees potentially have access to 35–47% of the average daily production of floral nectar in E. incrassata and actually harvest considerably less. These data show that E. incrassata flowers are adapted to restrict insect foragers despite their superficially unspecialized appearance. Eight forest and woodland eucalypts do not have a flower stage which excludes insects and the significance of this difference is discussed.  相似文献   

13.
Luis Navarro 《Biotropica》1999,31(4):618-625
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.  相似文献   

14.
Relationships between ornithophilous flowers and hummingbirds have been little studied in southern South America, where hummingbird species richness is low. We studied an ornithophilous flower assemblage and the hummingbird pollinators in a montane forest in southeastern Brazil. Twenty-three native hummingbird-pollinated plant species in 21 genera and 14 families were observed. Bromeliaceae, Fabaceae, Gesneriaceae, and Lobeliaceae are represented by more than one species within the assemblage. Flower shapes vary from narrow tube to bowl-shape, but tubular flowers prevail. The variety of flower shapes and sizes results in diverse pollen placement on the body parts of hummingbird visitors, although pollen is deposited mostly on the bill. Sugar concentration in nectar averages 22.1%, and nectar volume per flower averages 16.9 μl. The plant populations bloom for one month to year-round, and their flowering approaches the steady-state pattern. Four flower subsets may be defined within the assemblage, each subset related to the bill size and foraging habits of the most frequent bird visitor. Of the six species of hummingbirds recorded at the study site, four are common and largely resident. The four hummingbirds differ in bill size, body mass, and favoured foraging sites, attributes which reflect their favoured flower subsets. One hermit and one trochiline hummingbird share most of the flower species they use, these two birds being the major pollinators within the flower assemblage. This montane forest community may be viewed as medium-rich in ornithophilous flower species and poor in hummingbird species.  相似文献   

15.
Many species of honeyeaters and other nectar-feeding birds occur in most habitats in South Australia. They frequently feed on nectar of the same species of plants. A succession of species of plants provide nectar for birds throughout the year. Nectar is most abundant in winter and early spring and least abundant in summer and autumn. There is more nectar per flower and more flowers in winter and spring. Nectar is often depleted by honeyeaters, and sometimes other visitors (silvereyes, lorikeets and insects) between December and May. It is at times reduced to a level at which it is uneconomical for some species to exploit. There are seasonal movements of honeyeaters into areas of abundant nectar and out of these areas when nectar becomes scarce. Breeding coincides with peak abundance of nectar. Diversity of honeyeaters is probably maintained by an interaction of two types of competition, exploitation and interference. The larger species use the richest sources of nectar and aggressively exclude the smaller species (interference) whereas the smaller species can use poorer sources of nectar because their energy requirements are less (exploitation).  相似文献   

16.
Energy regulation by traplining hummingbirds   总被引:2,自引:0,他引:2  
1. A published model of constant diurnal energy accumulation by territorial hummingbirds does not accurately reflect the temporal distribution of feeding behaviour of traplining hummingbirds, Phaethornis longirostris (Long-Tailed Hermit Hummingbirds).
2. In an enclosure study, gross nectar intake by P . longirostris decreased through the day, mirroring nectar production rates in its natural food-flowers and mimicking its natural foraging patterns.
3. Using a simulation model, the energetic consequences of constant and decreasing net energy intake rates for traplining hummingbirds are compared.
4. Given natural patterns of nectar production, model birds with decreasing diurnal net intake rates met their energetic needs with fewer flowers than those with constant net intake, and spent less time foraging.
5. It is concluded that P . longirostris do not satisfy the physiological assumptions of the published model, and that in this way they are different from the territorial species on which the model has previously been tested.  相似文献   

17.
Interspecific competition can strongly influence community structure and limit the distribution and abundance of species. One of the main factors that determine hummingbird community structure is competition for food. The temporal and spatial distribution of nectar has a strong impact on hummingbird assemblages, shaping foraging niches according to hummingbird dominance and foraging strategy. We investigated whether body size and the degree of aggressive dominance influence feeding behaviour of hummingbirds in a temperate forest in northwestern Mexico (El Palmito, Mexico) when winter migrant hummingbirds are present in the community. First, we determined the dominance status of hummingbirds and evaluated the relationship between dominance and body mass, wing disc loading and migratory status. Secondly, we determined how hummingbird species used plant species differently. Thirdly, we examined whether the most dominant hummingbird species defended floral patches with more energy and/or with a larger number of flowers. At each flower patch, hummingbird species, number of hummingbird interactions, feeding time and number of flowers present were recorded. The total number of calories available within each floral patch was also determined. Our results demonstrate that the dominance hierarchy of 13 hummingbird species (migratory and resident) was correlated with body size but not wing disc loading, and that members of the hummingbird community showed a clear separation in resource use (by plant species). Hummingbirds at the top of the dominance hierarchy defended and fed on the best flower patches, defined by the quantity of calories available. Hence, the feeding behaviour of hummingbirds at El Palmito depends on the abundance of plant species used by hummingbirds and on the amount of energy available from each flower patch. Thus, hummingbird body size, aggressive dominance and defence of quality flower patches determines niche partitioning among species.  相似文献   

18.
It is normally thought that deep corolla tubes evolve when the plant's successful reproduction is contingent on having a corolla tube longer than the tongue of the flower's pollinators. Combining optimal foraging theory and quantitative genetics in a spatially explicit, individual-based model, we show that flowers with long corolla tubes can alternatively evolve because they promote resource partitioning among nectar feeders and increase the probability of conspecific pollen transfer. When there is competition for resources, long-tongued flower visitors feed preferentially at deep flowers and short-tongued visitors at shallow flowers. Both plant species thus benefit when the depths of their corollas are so different that each flower visitor specializes on one species. Resource competition can promote the evolution of deep corollas despite the presence of significant amounts of noise, such as deviations from optimal foraging behavior due to perceptual errors or temporal fluctuations in the relative abundance of competing pollinator species. Our results can explain the evolution of long corollas in a number of systems that do not conform to the traditional view.  相似文献   

19.
This field study shows that the number of flowers visited per bee per plant (Anchusa officinalis) increases with the instantaneous nectar level at the plant. Observations during the season showed that a bee visits more flowers per plant of given nectar level, the lower the overall mean nectar level in the study area. These results agree with predictions from a model based on the ‘marginal value theorem’, but with assumptions and constraints adapted for nectar-foraging bees. It suggests that bumblebees assess the nectar level at a plant by sampling one or a few flowers, which is possible because within-plant nectar volumes are correlated. The bees compare encountered gains to an optimal plant switching threshold equal to the overall mean nectar level and leave an unrewarding plant as soon as possible, but continue to visit the flowers on a rewarding plant. However, the bees leave before having visited all flowers due to a searching constraint. The bees’ response to plant nectar levels results in systematic flower visitation, because visitation to recently depleted flowers is reduced, which reduces the variation of the inter-visit time per flower. Systematic flower visitation implies that the overall mean encountered gain per flower is higher than the overall mean standing crop, as predicted by a model of systematic foraging. However, the sampling and searching constraints on the bees’ response to plant nectar levels increase the variation of the inter-visit time per flower, and thereby limit the degree of systematic flower visitation and the effect on the mean encountered gain.  相似文献   

20.
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.  相似文献   

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