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 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.     

Nectar utilization and pollination by Australian honeyeaters and insects visiting Calothamnus quadrifidus (Myrtaceae)

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.     

Pollinator visits to floral colour phases of Fuchsia excorticata

Abstract

Fuchsia excorticata is a gynodioecious tree (endemic to New Zealand) which is pollinated by honeyeater birds. Red, tubular flowers are common among bird-pollinated plants, and the tubular flowers of F. excorticata change colour from green to red. The purpose of the present study was to describe the timing of the colour change, dropping of the floral tube, and nectar production of F. excorticata and to determine how bellbirds (Anthornis melanura) and two introduced species of nectar robbers (Zosterops lateralis and Bombus sp.) respond to the different colour phases.

Floral tubes fell off about 11 days after anthesis in both sexes, with colour change occurring on about Day-4 for female trees and on about Day-5 for hermaphrodite trees. Green-phase hermaphrodite flowers produced significantly more nectar/day than did green-phase female flowers, while red-phase flowers did not produce nectar in either sex. All three floral visitors studied preferentially visited green-phase flowers and virtually ignored the nectarless flowers in the red phase. These results contrast with the general association between red, bird-pollinated flowers and the presence of a nectar reward. We suggest that the non-migratory habit of the New Zealand honeyeaters and the lack of native insect visitors to this species may account for this anomalous green-to-red colour change.     

How specialized is the plant–pollinator association between Eucalyptus globulus ssp. globulus and the swift parrot Lathamus discolor?

Abstract The swift parrot Lathamus discolor (Shaw) (Psittacidae) evolved from granivorous ancestors to become a specialized flower‐feeder in a monotypic genus. Its reproduction is dependent largely on flowers of Eucalyptus globulus Labill. ssp. globulus (Myrtaceae), the birds migrating to breed within the natural distribution of this tree. This paper investigates the extent to which this dependence of L. discolor on E. globulus is mirrored by dependence of the tree on the bird. It was found that L. discolor carried significantly more eucalypt pollen within 22 mm of its bill tip than did the New Holland honeyeater, Phylidonyris novaehollandiae (Latham) (Meliphagidae), and that pollen was concentrated on the regions of the head of L. discolor that consistently contact stigmas. Larger pollen loads on L. discolor can be attributed to it consuming both pollen and nectar, while honeyeaters take nectar only. The short thick bill of L. discolor necessitates regular stigmatic contact while the long slender bills of honeyeaters are unlikely to contact stigmas as often in these bowl‐shaped flowers. These factors suggest that L. discolor has a greater capacity to deposit pollen on stigmas of E. globulus than do honeyeaters. However, the characteristics of L. discolor that make it such an effective pollinator of E. globulus are also exhibited by a lorikeet (Psittacidae) that feeds on flowers of E. globulus. The association between E. globulus and L. discolor is therefore only moderately specialized because the flowers are also adapted to the more recently associated lorikeet and are almost certainly also pollinated by honeyeaters.     

Effects of flower removal on abundance and behaviour of honeyeaters in heathland near Sydney

Removal of almost all honeyeater nectar sources from a 5.6 ha area during February to July had no apparent effect on honeyeater nesting or total abundance. Behaviour in the experimental area was also generally unaffected except for the extent of nectar-feeding which was significantly reduced for the most commonly observed species. The honeyeaters must have compensated for the nectar removal by flying to nearby productive areas to feed on nectar. Nectar-removal had no significant effect on the species composition of resident honeyeaters but did affect the species composition of all observed honeyeaters. Nectar removal resulted in a decrease in the abundance of the large and dominant Little Wattlebirds (Anthochaera chrysoptera) and an increase in the abundance of the other, smaller species.     

Ecomorphological constraints imposed by the kidney component measurements in honeyeater birds inhabiting different environments

The histological renal anatomy of 10 species of honeyeaters was examined quantitatively, using stereology. The kidneys of five species of predominantly wet zone inhabiting birds: the western spinebill Acanthorhynchus superciliosus , White-checked honeyeater Phylidonyris nigra . New Holland honeyeater Phylidonyris novaehollandiae , little wattlebird anthochaera chrysoptera and red wattlebird Anthochaera carunculata , were compared to five predominantly arid zone inhabiting birds: the grey-fronted honeyeater Meliphaga plumula , white-plumed honeyeater Meliphaga penicillata , white-fronted honeyeater Phylidonyris albifrons , spiny-checked honeyeater Acanthogenys rufogularis and yellow-throated miner Manorina flarigula. The kidneys were asymmetrical, with the left kidney being larger than the right kidney. Kidney mass was directly proportional to body mass (coefficient of correlation, r =+0·95), as was kidney volume to kidney mass (r =+1·0). Wet zone honeyeaters generally and a higher percentage and absolute volume of renal cortex, whilst arid zone honeyeaters generally had a significantly higher percentage and absolute volume of renal medulla. There were few differences between species, in either the percentage or absolute volume or luminal surface area of nephron components within the cortex. Within the medulla, wet zone honeyeaters generally had a higher percentage and absolute luminal surface area of collecting ducts, whilst arid zone honeyeaters had a higher percentage and absolute surface area of capillaries. This may be due to factors such as variations in diet and climate between habitats resulting in differences in honeyeater renal morphology which enable arid zone birds to conserve water and wet zone birds to conserve ions more efficiently.     

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.     

Territoriality and changes in resource use by sunbirds at Leonotis leonurus (Labiatae)

Summary Leonotis leonurus was studied with the aim of characterising nectar quality and quantity, and of monitoring the sunbirds' response to the resource. The flowers produced high quality nectar in low quantities. Nectar quality remained constant during the day and averaged 23.4% sucrose equivalents (an energy content of 3.8j/l). Nectar production was highest during the morning when flow rates reached a peak of 3.3 l/h. L. leonurus was visited by four species of sunbirds. In 1977 there were 7.8 visits/h to the flowers. The clump was defended by a single Olive Sunbird (Nectarinia olivacea) who defended the flowers intra- and interspecifically. On one day the territorial bird obtained approximately 57.1 kJ from the Leonotis flowers and expended 5.4 kJ on defence and 13.6 kJ overall while on the territory. Territorial behaviour and intruder pressure was a function of resource availability. When Leonotis nectar levels were high intruder pressure was intense and the territorial bird employed long chases as a defence behaviour. When nectar levels desreased intruder pressure was lower, the territorial bird chased less and employed vocalisations as a relatively inexpensive form of territorial behaviour.In 1978 there was an abundance of nectar sources in the study area due to the late onset of the dry season. Sunbirds were not territorial at Leonotis and visiting rates were lower than in 1977, averaging only 1.7 visits per hour. Instead the sunbirds concentrated on contemporaneously flowering plant species, particularly Halleria lucida.     

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.     

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.     

Morphological relationships of passerine birds from Australia and New Guinea in relation to their diets

Beak, wing, leg and intestinal lengths, and gizzard widths, were all significantly related to body mass in 51 honeyeater species from Australia, 48 honeyeater species from New Guinea and 31 purely insectivorous passerine bird species from Australia. The nectar-feeding honeyeaters had smaller gizzards and intestines than wholly insectivorous birds of comparable size, although their wing and leg lengths did not differ; New Guinean and Australian honeyeaters were similar in these respects. Overall, honeyeaters had longer beaks than pure insectivores. Among Australian honeyeaters, those genera consuming more nectar than insects had longer beaks than the less nectarivorous, more insectivorous genera. Indeed, the latter group had beaks comparable in length to wholly insectivorous birds. All morphological differences revealed were attributable to known differences in diet.     

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.     

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).     

Seasonal changes in the abundance and body condition of honeyeaters (Meliphagidae) in response to inflorescence and nectar availability in the New England National Park,New South Wales

Seasonal changes in numbers and body condition of honeyeaters were examined over 2 years in the New England National Park, New South Wales, Australia. Nectar availability measured as inflorescence density and nectar production were also recorded. In the main study site, the abundance of six of the seven most common species of honeyeater was significantly correlated with inflorescence density. However, honeyeater numbers were not significantly correlated with daily energy productivity. Unlike most honeyeater communities, the New England community was dominated numerically by the Eastern Spinebill, the smallest of the most common species present. Among the most common species, the duration of time a species was recorded in the area was negatively, but not significantly, correlated with its size. All species examined showed similar seasonal changes in body condition, with the highest bodyweights and fat deposits recorded in winter.     

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.     

Quantifying the canopy nectar resource and the impact of logging and climate in spotted gum Corymbia maculata forests

Nectar in tall forest canopies is a significant, but unquantified resource for Australian fauna. We investigated the impact of logging on nectar production in the canopy of spotted gum Corymbia maculata in southern New South Wales. In addition, we quantified the magnitude of canopy nectar production and how this varied with climate over 2 years. In 2005 flowers were bagged on large and small trees in replicate recently logged, regrowth and mature forest. Neither logging history nor tree size significantly affected overnight nectar production per flower, although there was a significant interaction. When nectar production was scaled up to the forest stand (incorporating flower and tree density) mature forest produced almost 10 times as much sugar per ha as recently logged forest, with regrowth being intermediate. Under current forest practices at the compartment scale, the difference between mature forest and recently logged forest was reduced to a factor of two times. One distinctive characteristic of C. maculata nectar in 2005 was its high sugar content (40–60%) compared with the concentrations measured in 2003 (～18%). Nectar was only slightly depleted in unbagged flowers in 2005 when flowering was unusually extensive. We estimated that, on average, mature spotted gum forest produced a vast resource of nectar overnight: 35 000 Kj ha^?1. Flowers measured in 2003 provided a strong contrast with only occasional stands of trees flowering, much less sugar per flower early in the morning and unmeasurable quantities by mid‐morning, indicating that nectar was limiting. Measurements at sites in 2003 indicated that regrowth sites could be more productive than mature forest; however, few sites were measured. We suggest that management should focus mitigations on poor flowering years when the nectar resource is limiting. Models of nectar production collated over both years, using climate and site variables, indicated nectar volumes and sugar concentration respond differently to environmental conditions. Predicting the nectar resource, which is made up of both components, was most consistently related to recent conditions that were unfavourable to foliage production.     

The seasonal abundance and foraging behaviour of honeyeaters and their potential role in the pollination of Banksia menziesii

Seasonal changes in the abundances of five species of honey-eaters were assessed in relation to the flowering phenology of Banksia menziesii in banksia woodland near Perth, Western Australia. The total number of honeyeaters was significantly correlated to the number of inflorescences ofB. menziesii. New Holland Honeyeaters. Brown Honeyeaters and Western Spinebills were present throughout the year, whereas the larger honeyeaters (Red and Little Wattlebirds) were observed when B. menziesii was flowering. The foraging behaviours of the Little Wattlebirds, New Holland Honeyeaters, Brown Honeyeaters and Western Spinebills were similar and all were likely to effect pollination of B. menziesii florets. Differences in their foraging position at an inflorescence, number and direction of foraging probes, and the time spent at an inflorescence were minimal. Western Spinebills used inflorescences lower in the tree crown than the other species. Movements to inflorescences on different plants were inversely related to honeyeater size. Smaller honeyeaters were frequently chased from inflorescences by the larger species, increasing the proportion of distant foraging movements made by the smaller species. Also these interrupted visits were of shorter duration than uninterrupted visits. Visitation by smaller species, especially the Brown Honeyeater, may result in more cross pollinations although the effect on the reproductive success of B, menziesii is unknown.     

Coexistence and Habitat Preference of Two Honeyeaters and a Sunbird on Lombok,Indonesia

On the island of Lombok, Indonesia, three nectarivorous birds partially coexist: the two closely related and very similar Lichmera honeyeaters and a sunbird (Cinnyris jugularis). We investigated how these species segregated ecologically in areas where they coexisted by evaluating foraging visits and aggressive interactions at rich and poor nectar resources in different habitats (forest and open areas) along an altitudinal gradient (800–1600 m asl). The two honeyeaters were partially segregated by altitude. In the zone of overlap, Lichmera limbata dominated the richest forest resources, while Lichmera lombokia dominated the richest resources in open land. The sunbird, C. jugularis, was only observed in open habitats and mostly at poor resource patches. In the three‐species community in open habitats a dominance hierarchy was apparent with L. lombokia as the superior species and C. jugularis as the inferior species. Studies on how segregated species interact in their transition zone can help us to understand interactions between otherwise ecologically segregated species.     

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.