Honeyeater population changes in relation to food availability in the Jarrah forest of Western Australia

The numbers of honeyeaters present at particular sites in the Jarrah forest varied significantly from month to month, with peak abundance occurring between May and September. Numbers also varied from site to site, depending upon the major plant species present. Honeyeater abundance was not limited by arthropod availability, but in many instances was closely correlated with the availability of nectar, particularly that produced by Dryandra sessilis. Large honeyeaters, such as Anthochaera chrysoptera and Phylidonyris novaehollandiae, were generally most abundant at times and sites of greatest nectar production. Small honeyeaters, such as Acanthorhynchus superciliosis, were never abundant but were present for most of the year. The production of nectar between October and December was such that more honeyeaters could have been supported than were actually present. Low numbers at these times can be explained in terms of reduced foraging efficiency that would have resulted from more widely dispersed flowers, and the possible availability of more rewarding nectar resources at other sites.     

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.     

Mutualistic interdependence between mistletoes (Amyema quandang), and spiny-cheeked honeyeaters and mistletoebirds in an arid woodland

The mutualism involving mistletoes (Amyema quandangj, spiny-cheeked honeyeaters (Acan-thagenys rufogularis) and mistletoebirds (Dicaeum hirundinaceum) was studied in arid woodland in South Australia between 1980 and 1984. Plants and birds were locally interdependent: mistletoes supplied a continuous resource of fruits or nectar that sustained permanent populations of pollinators (honeyeaters) and dispersers (honeyeaters and mistletoebirds). The reproductive phenology of Amyema quandang was central to the interactions. Amyema quandang flowered in winter and annual fruit crops overlapped so that ripe fruit was continuously available. Spiny-cheeked honeyeaters obtained most of their energy requirements from mistletoe nectar in winter and mistletoe fruit in summer. Higher honeyeater densities were sustained by flowering in winter. Mistletoebirds were present in low density throughout the year and subsisted on a diet of mistletoe fruit and a few insects. The reproductive strategy of A. quandang probably evolved in response to the pollination and dispersal service provided by honeyeaters in inland Australia. Neither spiny-cheeked honeyeaters nor mistletoebirds have adaptations resulting from evolutionary interactions with A. quandang. The high specificity of their mutualism is a result of: (i) the abundance of A. quandang in relation to other nectar and fruit producing plants in the community: (ii) the year-round production by A. quandang of the primary source of fruit or nectar for honeyeaters and mistletoebirds: (iii) the facultative specialization of both birds on A. quandang; and (iv) the reluctance or inability of other frugivorous birds in the community to consume A. quandang fruit.     

The diet of the New Holland honeyeater,Phylidonyris novaehollandiae

New Holland honeyeaters collect nectar, manna or honeydew for energy and hawk small flying insects for protein. The insects taken were usually Diptera and Hymenoptera weighing 0.7 mg dry weight or less. Net rates of energy gain from hawking small flying insects were usually less than 20 J min^?1 and sometimes negative and insufficient to meet the bird's daily energy requirements. Those from feeding on nectar, manna or honeydew were usually above 40J min^?1 and often above 400J min^?1 at dawn and the birds depended on these carbohydrates for energy. Nectar, manna and honeydew contained negligible amounts of protein, and the birds used small flying insects as sources of protein, and presumably other nutrients. Given that carbohydrate resources supply better rates of energy gain than insects. New Holland honeyeaters should collect their energy requirements from carbohydrates and only collect sufficient insects to satisfy their protein requirements. Estimates of the food intakes of both non-breeding and breedig birds showed that they did this. Non-breeding New Holland honeyeaters collected from 72 to 125 (mean 92) kJ of carbohydrates per day and 17 to 58 (mean 31) mg of protein per day. These meet the daily energy (75 kJ) and protein (20 mg) requirements of the birds. Breedig birds collected more carbohydrates and more insects, but in proportion to their increased energy and protein requirements respectively. New Holland honeyeaters are probably limited by their ability to meet their energy requirements from nectar, manna or honeydew and not by insects. Non-breeding birds collected their protein requirements in about 10 min of insect-feeding, but spent from 33 to 90% of the day collecting carbohydrates to meet their energy requirements. The maintenance requirement of 20 mg of protein per day for New Holland honeyeaters is about 25% of that estimated from standard equations for a bird of the same size. This low level may have evolved in response to low energy availability.     

Partitioning of nectar sources in an Australian honeyeater community

Feeding by honeyeaters was found to maintain nectar at low levels at three sites studied on Kangaroo Island in May-June 1978. The productivity of nectar at a site and position in a dominance hierarchy appeared to determine which bird species used each site. Correa was the main nectar source in the poorest area and produced 0.05 kJ m^?2 per day. The small eastern spinebill was the most abundant honeyeater. The purple-gaped honeyeater also occurred but fed mostly on honeydew. The medium sized New Holland honeyeater was common and territorial in the second area, where Banksia marginata and B. ornata inflorescences and Adenanthos flowers produced 0.7 kJ m^?2 of nectar per day. Spinebills and crescent honeyeaters also visited flowers and were sometimes chased by New Holland honeyeaters. The richest site was a flowering Eucalyptus cosmophylla tree (5.1 kJ m^?2 of nectar per day). A red wattlebird, the largest honeyeater, held a territory in part of this tree and chased other honeyeaters from the territory. New Holland, crescent and purple-gaped honeyeaters fed on flowers in other parts of the tree. The spinebill was absent. We conclude that nectar was partitioned along a spectrum of rich to poor sources. Larger more aggressive species used and sometimes defended the richest sources while the smaller birds used the poorer sources.     

Honeyeater foraging: A test of optimal foraging theory

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.     

Consequences of differences in body mass,wing length and leg morphology for nectar-feeding birds

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.     

The exploitation of floral nectar in Eucalyptus incrassata by honeyeaters and honeybees

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.     

Contrasting movement patterns of nectar-collecting and pollen-collecting bumble bees (Bombus terricola) on fireweed (Chamaenerion angustifolium) inflorescences

Abstract. 1. Movements of nectar and pollen-foraging bumble bees on inflorescences of Chamaenerion angustifolium (L.) J. Holub (fireweed or rosebay willow herb) were compared with predictions based on reward distributions and optimality principles. Observations suggest that nectar and pollen-gathering bumble bees behave according to the same set of reward maximization criteria when foraging from flowers of this species.
2. Both kinds of foragers matched their arrival points with the vertical positions on inflorescences in which the densities of their respective food resources were greatest. For nectar-foragers, this point was located at the lowest tier of flowers, whereas for pollen-foragers it was found in the middle of the inflorescences. Nectar and pollen-foraging bees both moved upward on inflorescences following gradients from high to low reward availability.
3. Nectar-foragers responded to decreases in inflorescence size over the season by reducing the number of flower visits made on each raceme. Number of flowers visited by pollen-foragers was low throughout and reflected the scarcity of male-phase flowers on racemes. Flower revisitation rates were low for both kinds of workers, but were slightly higher for those collecting pollen.     

Seasonal changes in a honeyeater assemblage in Banksia woodland near Perth,Western Australia

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.     

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 birds and the bees: pollinator behaviour and variation in the mating system of the rare shrub Grevillea macleayana

Background and Aims

In Australia, honey-bees have invaded systems that evolved without social insect pollinators, where many plants are adapted to vertebrate pollination. Behavioural differences between pollinators are likely to influence mating patterns, but few studies have examined this empirically in long-lived, woody, perennials. It was shown previously that outcrossing rates in Grevillea macleayana vary among populations. Here tests were conducted to determine whether the behaviour of birds and honey-bees differed between a population previously found to be highly outcrossed and two inbreeding populations.

Methods

Visit frequencies and movement patterns of the visitors to inflorescences at three sites over two seasons were compared. A caging experiment was used to test the effects of excluding birds on pollen removal from newly opened flowers and on pollen deposition on stigmas that had been washed clean.

Key Results

Honey-bees were the most frequent visitors overall, but honeyeaters were more frequent visitors in the population previously found to have a high outcrossing rate than they were in either of the other populations. More visits by honeyeaters were from distant plants. Pollen removal did not vary greatly among sites, and was not affected by bird exclusion; however, more pollen was deposited on the stigmas of cleaned pollen presenters in the population previously observed to be highly outcrossing than in the other two. This high level of pollen deposition was reduced by experimental bird exclusion.

Conclusions

Honey-bees were the most frequent visitors, by an order of magnitude, and excluding vertebrates revealed that bees were removing most of the pollen but deposited fewer pollen grains on stigmas. Birds were more frequent visitors at the site previously found to be outcrossing than the other two sites, and they moved further between plants and visited fewer inflorescences on a plant during a foraging bout than bees did. These characteristics of bird visits to G. macleayana would be sufficient to produce significant variation in outcrossing rates among sites.Key words: Grevillea macleayana, Apis mellifera, honey-bees, honeyeaters, pollinator behaviour, pollen removal, pollen deposition, outcrossing rate     

Interspecific competition in Australian honeyeaters—depletion of common resources

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).     

Differences in pollinator effectiveness of birds and insects visiting Banksia menziesii (Proteaceae)

Summary The effectiveness of nectarivorous birds, introduced honey bees and staphylined beetles as pollinators of Banksia menziesii was assessed. Staphylinids removed substantial amounts of pollen but did not deposit any onto stigmata. Abundance of beetles on inflorescences was related to the mean number of florets opening per day. Honey bees collecting pollen were more likely to effect pollination than those collecting nectar which only contacted stigmata when arriving or leaving an inflorescence. Nectar-foraging birds probed between florets 10.2±0.8 (±SE) times, contacting 8–16 stigmata during each probe. Bees visited inflorescences ten times more frequently than birds although they deposited only 25% of the pollen that birds did on stigmata. Fruit set was ten times greater on inflorescences visited by birds than on inflorescences visited by bees. Bees were capable of removing as much pollen as birds but, because of direct pollen transfer to birds when florets opened during foraging, actual removal was probably much less. Selection for floret opening during nectar foraging by birds may have resulted from pollen removal by non-pollinating animals, such as staphylinids.     

Bills and tongues of nectar-feeding birds: A review of morphology,function and performance,with intercontinental comparisons

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.     

Response of avian nectarivores to the flowering of Aloe marlothii : a nectar oasis during dry South African winters

In southern Africa, Aloe marlothii flowers during the dry winter season and offers copious dilute nectar to a variety of birds. Avian abundance and community composition were monitored at an A. marlothii forest at Suikerbosrand Nature Reserve, South Africa. Sampling occurred during two summer months (February–March) when no flowers were present, and six months (May–October) that spanned the winter flowering. We hypothesized that an influx of occasional nectarivores to the A. marlothii forest during flowering would lead to significant changes in the avian community. Overall bird abundance increased 2–3 fold at the peak of nectar availability (August). We recorded 38 bird species, of 83 species detected during transects, feeding on A. marlothii nectar; this diverse assemblage of birds belonged to 19 families, including Lybiidae, Coliidae, Pycnonotidae, Sylviidae, Cisticolidae, Muscicapidae, Sturnidae, Ploceidae and Fringillidae. Surprisingly, only two species of sunbird (Nectariniidae) were observed feeding on A. marlothii nectar, and both occurred in low abundance. We predicted that competition for nectar resources would be high, but few aggressive inter- and intra-specific interactions occurred between birds while feeding on inflorescences. During peak flowering, insect feeders (insectivores, omnivores, nectarivores) fed on nectar during the cold morning when insect activity was low, whilst non-insect feeders (frugivores and granivores) fed on nectar in the middle of the day. Our study highlights the importance of A. marlothii nectar as a seasonal food and water source for a diverse assemblage of occasional nectarivores.     

Effectiveness of nectarivorous birds and honeybees as pollinators of Banksia spinulosa (Proteaceae)

Abstract The effectiveness of nectarivorous birds and honeybees (Apis mellifera) as pollinators of Banksia spinulosa (Proteaceae) was investigated. Birds visited inflorescences in the early, mid and late flowering seasons. In contrast, honeybees visited only on days in the late flowering period when maximum temperatures exceeded 15°C. Self pollen remained on pollen presenters of flowers for up to 5 days in the early and mid periods. In the late period, when honeybees visited inflorescences, self pollen was removed within 2 days. Pollen removal was similar for caged (birds excluded) and open inflorescences in the late period, indicating that most pollen was removed by honeybees. In the early and mid periods, honeyeaters pollinated 22% and 27% of flowers on open inflorescences, respectively. In the late period, when both birds and bees visited inflorescences, 64–73% of flowers on open inflorescences were pollinated. Foraging by honeybees resulted in pollen deposition as 38% of flowers on caged inflorescences were pollinated. Throughout the flowering season a similar number of pollen grains was deposited per stigma. There were 3.0–3.7 pollen grains per stigma on open inflorescences in the late period, although only 2.0 grains per stigma on caged inflorescences. In the early and mid periods, fewer caged than open inflorescences produced fruits, indicating the importance of honeyeaters to reproductive success at these times. In contrast, in the late period when honeybees visited inflorescences, fruit-set was similar on caged and open inflorescences. Overall, these results indicate that honeybees were effective pollinators of B. spinulosa.     

Seasonal variation in honeyeater foraging behaviour,infloresence abundance and fruit set in Banksia spinulosa (Proteaceae)

Seasonal variation in the foraging behaviour of honeyeaters and the production of fruit were examined in relation to the flowering intensity of Banksia spinulosa over two flowering seasons. The abundance of inflorescences was greater in the mid than in the early and late periods of the flowering season. In the mid period, many plants were blooming and each plant had many flowering inflorescences. Inflorescences received most visits by honeyeaters in the early flowering period; the visitation rate declining as flowering progressed. Eastern spinebills were the most common floral visitors at all times during the season. The number of foraging probes made at inflorescences by eastern spinebills did not differ throughout the season. Foraging movements between inflorescences on the same plant were more frequent in the mid period than in the early and late periods. Long distance movements between plants (more than 10 m apart) were promoted by aggressive interactions between honeyeaters. Inflorescences flowering in the late period were less likely to develop follicles because there were fewer visits by birds and/or because resources had been allocated to inflorescences pollinated earlier in the season. The number of follicles produced per infructescence did not differ between flowering periods. Overall, the number of inflorescences produced per plant, the number of visits received per inflorescence and the proportion of inflorescences that developed follicles were greater in 1987 than in 1988.     

Honeyeaters and the New Zealand forest flora: The utilisation and profitability of small flowers

New Zealand flowers are frequently considered unspecialised allowing easy access to pollen and nectar by a wide range of visitors. Most conform with a syndrome of insect pollination (entomophily). Pollination of forest flowers by birds has been described for a range of species whose flowers are morphologically ornithophilous. On Kapiti Island and Little Barrier Island, all three species of New Zealand honeyeaters have been described feeding on flowers currently assumed to be entomophilous or where the pollination system is unknown. The persistence and regularity of visits suggests that the birds are obtaining suitable rewards in the form of nectar and could be serving as pollinators. We measured the nectar energetic value from flowers of three ornithophilous and five entomophilous species. Nectar production over 24 hours was highest in ornithophilous species, but the standing crop of nectar overlapped—ornithophilous species: Metrosideros fulgens (standing crop 6.6 J), Metrosideros excelsa (22 J), and Fuchsia excorticata (1.8 J); and entomophilous: Pittosporum crassifolium (23 J), Pseudopanax arboreus (1.5 J), Dysoxylum spectabile (3.7 early flowers -6.7 J late flowers), Pittosporum eugenioides (2.7 J) and Geniostoma rupestre (1.8 J). The entomophilous species present the flowers in aggregation and as result birds can visit a large number flowers per minute. We found that the average estimated nectar consumption rate for all the entomophilous species except G. rupestre was enough to sustain the two smaller New Zealand honeyeaters (hihi energy requirements= 0.12 kJ min(-1), median energy obtained: 0.16 kJ min(-1) D. spectabile—0.57 kJ min(-1) P. crassifolium); bellbird energy requirements = 0.10 kJ min(-1), median energy obtained: 0.14 kJ min(-1) D. spectabile—0.68 kJ min(-1) P. crassifolium). However, we estimate that if the birds are able to selectively forage on the flowers with most nectar, the energetic returns of all species may be sufficient for hihi and bellbird (hihi: 0.18 kJ min(-1) G. rupestre—0.93 kJ min(-1); P. crassifolium; bellbird: 0.12 kJ min(-1) G. rupestre 1.11 kJ min(-1) P. crassifolium). If tui (energy requirements: 0.25 kJ min(-1), forages randomly, only P. crassifolium (0.80 kJ min(- 1)) and D. spectabile late in the season (0.30 kJ min(-1)) provide sufficient returns, but if selective, P. arboreus (0.45 kJ min(-1)) may also suffice. We suggest that because (a) the nectar produced by entomophilous flowers provides sufficient energy to sustain the energetic requirements of birds, and (b) these plants flower in the cooler months when insect activity is reduced, birds might have played a wider role in pollination than previously considered. This finding is of particular importance because the abundance of New Zealand honeyeaters on the mainland has decreased considerably since human colonisation and this could be affecting forest regeneration.     

Flowering phenology,seed set and bird pollination of five Western Australian Banksia species

Flowering phenology and seed set characteristics of five species of Banksia were studied in relation to the nectarivorous birds which feed at their inflorescences. Within the Banksia woodland at the study site near Perth, the flowering seasons of the Banksia species were sequential and only slightly overlapping, providing a year-round nectar source. Although honeyeaters visited alt five species, seed set was very low in each case. Caging experiments indicated that, in B. attenuata at least, alternative pollinators may play a more important role in pollination than do nectar-feeding birds. It is suggested that non-avian pollinators, predatory insects, and characteristics of the breeding system may also have been important in the evolution of the observed flowering phenology and patterns of seed set.