New Holland Honeyeaters Research Articles

How do honeyeaters drink nectar?

We investigated the kinematics and biomechanics of nectar feeding in five species of honeyeater (Phylidonyris novaehollandiae, Acanthagenys rufogularis, Ptilotula penicillata, Certhionyx variegatus, Manorina flavigula). There is abundant information on honeyeater foraging behaviors and ecological relationships with plants, but there has never been an examination of their nectar-feeding from kinematic and biomechanical perspectives. We analyzed high-speed video of feeding in captive individuals to describe the kinematics of their nectar feeding, with specific focus on describing tongue movements and bill-tongue coordination, and to characterize the mechanism of nectar uptake in the tongue. We found clear interspecific variation in kinematics and tongue filling mechanics. Species varied in lick frequency, tongue velocity, and protrusion and retraction duration, which, in some cases, are relevant for differences in tongue filling mechanisms. We found support for the use of capillary filling in Certhionyx variegatus only. By contrast, Phylidonyris novaehollandiae, Acanthagenys rufogularis, Ptilotula penicillata, and Manorina flavigula employed a modified version of the expansive filling mechanism seen in hummingbirds, as there was dorsoventral expansion of the tongue body, even the portions that remain outside the nectar, once the tongue tip entered the nectar. All species use fluid trapping in the distal fimbriated portion of the tongue, which supports previous hypotheses describing the honeyeater tongue as a "paintbrush."

Distress responses during handling in urban and exurban bandiny, the New Holland honeyeater (Phylidonyris novaehollandiae), in southwestern Western Australia

Context Birds appear to be better suited than mammals or reptiles to adapt to fast-changing landscapes because of their greater mobility; however, the behavioural effects of urbanisation on birds in Australia remain broadly unexplored. Aims This study aimed to investigate the effects of urbanisation on behavioural responses exhibited by a common and widespread meliphagid, the bandiny or New Holland honeyeater (Phylidonyris novaehollandiae) while undergoing standard bird banding processes. Methods Five non-invasive techniques (alarm calling, wriggling, biting, breathing rate, and tonic immobility) were explored for efficacy in identifying underlying differences in distress arising from mist-netting at urban and exurban localities in southwestern Western Australia. Key results Breathing rate was the most important variable for identifying differences in post-capture distress response. The breathing rate of urban bandiny following capture was lower than those of exurban areas. All other parameters proved suboptimal for detecting differential behavioural responses to handling between urban and exurban populations, despite having been successfully used in other international studies. Conclusions We suggest that urban honeyeaters exhibit lower mean breathing rates due to chronic overstimulation in response to urban lifestyles and are not able to further elevate this behaviour in response to stressful stimuli. The failure of other approaches explored highlights the need to apply ecosystem-appropriate methods for investigating urbanisation within an Australian context. Implications Our results suggest that behavioural approaches to quantifying avian stress developed internationally require additional consideration when applied to the ecosystems of Australia, whose birds are evolved to accommodate a different regime of seasonality that has shaped them behaviourally and morphologically.

Restored and remnant Banksia woodlands elicit different foraging behavior in avian pollinators.

Pollinators and the pollination services they provide are critical for seed set and self‐sustainability of most flowering plants. Despite this, pollinators are rarely assessed in restored plant communities, where their services are largely assumed to re‐establish. Bird–pollinator richness, foraging, and interaction behavior were compared between natural and restored Banksia woodland sites in Western Australia to assess their re‐establishment in restored sites. These parameters were measured for natural communities of varying size and degree of fragmentation, and restored plant communities of high and low complexity for three years, in the summer and winter flowering of Banksia attenuata and B. menziesii, respectively. Bird visitor communities varied in composition, richness, foraging movement distances, and aggression among sites. Bird richness and abundance were lowest in fragmented remnants. Differences in the composition were associated with the size and degree of fragmentation in natural sites, but this did not differ between seasons. Restored sites and their adjacent natural sites had similar species composition, suggesting proximity supports pollinator re‐establishment. Pollinator foraging movements were influenced by the territorial behavior of different species. Using a network analysis approach, we found foraging behavior varied, with more frequent aggressive chases observed in restored sites, resulting in more movements out of the survey areas, than observed in natural sites. Aggressors were larger‐bodied Western Wattlebirds (Anthochaera chrysoptera) and New Holland Honeyeaters (Phylidonyris novaehollandiae) that dominated nectar resources, particularly in winter. Restored sites had re‐established pollination services, albeit with clear differences, as the degree of variability in the composition and behavior of bird pollinators for Banksias in the natural sites created a broad completion target against which restored sites were assessed. The abundance, diversity, and behavior of pollinator services to remnant and restored Banksia woodland sites were impacted by the size and degree of fragmentation, which in turn influenced bird–pollinator composition, and were further influenced by seasonal changes between summer and winter. Consideration of the spatial and temporal landscape context of restored sites, along with plant community diversity, is needed to ensure the maintenance of the effective movement of pollinators between natural remnant woodlands and restored sites.

Morphological and molecular description of a new species of Isospora (Apicomplexa) from a New Holland honeyeater (Phylidonyris novaehollandiae).

A new Isospora species is described from New Holland honeyeaters (Phylidonyris novaehollandiae). Sporulated oocysts (n=25) were characterised as subspheroidal, 29-32×28-31 (29.8×29.4); length/width (L/W) ratio 1.01-1.02 (1.01). Wall bi-layered, 1.3-1.6 (1.5) thick, outer layer smooth, c.2/3 of total thickness. Micropyle and oocyst residuum absent, but usually two polar granules are present. Sporocysts (n=25) ovoidal, 18-19×12-14 (18.4×12.3); L/W ratio 1.42-1.53 (1.50). Stieda body present, flattened, c.0.5 deep × 2.5 wide; sub-Stieda present, rounded, c.2.5 deep × 3.5 wide; para-Stieda body absent; sporocyst residuum present, usually a distinctly irregular body consisting of numerous small granules that appear to be membrane-bound. Sporozoites vermiform, with robust anterior and posterior refractile bodies. Molecular characterization was conducted at the 18S and 28S ribosomal RNA and the mitochondrial (mt) cytochrome oxidase (COI) loci. Phylogenetic analysis of genomic 18S and mt COI sequences indicated that Isospora phylidonyrisae n. sp. was genetically similar to Isospora coronoideae, isolated from an Australian raven (Corvus coronoides) in Western Australia, with a 99.3% and 98.4% homology, respectively. The 28S rRNA sequence was most similar to Isospora anthochaerae (KF766053) and Isospora manorinae (KT224381), both with a 98.2% genetic similarity. Based on morphological and genetic data, this isolate is a new species of Isospora, which is named Isospora phylidonyrisae n. sp. after its host.

Speedy revelations: how alarm calls can convey rapid, reliable information about urgent danger

In the perpetual struggle between high-speed predators and their prey, individuals need to react in the blink of an eye to avoid capture. Alarm calls that warn of danger therefore need to do so sufficiently rapidly that listeners can escape in time. Paradoxically, many species produce more elements in their alarm calls when signalling about more immediate danger, thereby increasing the reliability of transmission of critical information but taking longer to convey the urgent message. We found that New Holland honeyeaters, Phylidonyris novaehollandiae, incorporated more elements in alarm calls given to more dangerous predators, but video analysis revealed that listeners responded in 100 ms, after only the first element. Consistent with this rapid response, the acoustic structure of the first element varied according to the danger, and playbacks confirmed that birds need hear only the first element to assess risk. However, birds hid for longer and were more likely to flee, after calls with more elements. The dual mechanisms of varying both element structure and number may provide a widespread solution to signalling rapidly and reliably about immediate danger.

Look at the time: diel variation in the flight initiation distance of a nectarivorous bird

When approached by a predator, foraging animals should adjust escape behavior based on their current energetic status to minimize opportunity costs (foraging time) and maximize the chance of escape. Animals should therefore be more reluctant to leave a foraging patch when their energy reserves are low and should flee using low-energy escape tactics, but be quicker to flee and use the best escape tactic when satiated. Nectarivorous birds have little capacity for energy storage; thus, they should be particularly sensitive to variation in nutritional status throughout the day. We tested whether New Holland honeyeaters (Phylidonyris novaehollandiae) alter their antipredator behavior by testing whether flight initiation distance (FID—the distance at which an individual flees from an approaching predator) and escape tactic (hopping or flying) vary over the course of the day. Honeyeater FID was shortest in the morning, following the overnight fast, and longest at midday. In the evening, though FID tended to shorten, responses were more variable overall, likely due to individual variation in foraging success. In the morning, honeyeaters were equally likely to hop or fly when approached, but increasingly favored flight as the day progressed. These results suggest a strong link between individual energetic condition and antipredator responses. We suggest that time of day is a useful and important variable influencing escape behavior, the potential effects of which should be more consistently considered as a covariate in the widely used FID protocol. Diel variation in internal and external conditions can alter behavioral responses. In this study, we observed antipredator response behaviors in foraging New Holland honeyeaters, small passerines sensitive to changes in energy status throughout the day. We found that honeyeaters allowed researchers to approach more closely in the morning and evening before fleeing and escaped by flying more often than hopping as the day progressed. Our results suggest that antipredator behaviors are sensitive to current energy status and impending energy needs. In addition, we show that time of day is an important consideration in the experimental design and statistical analysis of behavior studies.

Personal information about danger trumps social information from avian alarm calls.

Information about predators can mean the difference between life and death, but prey face the challenge of integrating personal information about predators with social information from the alarm calls of others. This challenge might even affect the structure of interspecific information networks: species vary in response to alarm calls, potentially because different foraging ecologies constrain the acquisition of personal information. However, the hypothesis that constrained personal information explains a greater response to alarm calls has not been experimentally tested. We used a within-species test to compare the antipredator responses of New Holland honeyeaters, Phylidonyris novaehollandiae, during contrasting foraging behaviour. Compared with perched birds, which hawk for insects and have a broad view, those foraging on flowers were slower to spot gliding model predators, showing that foraging behaviour can affect predator detection. Furthermore, nectar-foraging birds were more likely to flee to alarm call playbacks. Birds also assessed social information relevance: more distant calls, and those from another species, prompted fewer flights and slower reaction times. Overall, birds made flexible decisions about danger by integrating personal and social information, while weighing information relevance. These findings support the idea that a strategic balance of personal and social information could affect community function.

Do avian ticks (Ixodes hirsti) influence host phenotype in New Holland honeyeaters (Phylidonyris novaehollandiae)?

ABSTRACTParasites can impact on host phenotype, causing adverse population changes. In avian populations, increased feather asymmetry as a consequence of parasitism is one example of parasites impacting host phenotype. Feather asymmetry can be important in sexual selection, reproductive success and survival, but relationships between feather asymmetry and measures of survival remain unclear. Previous studies have demonstrated these relationships in avian populations through experimental manipulation of parasite loads, but few studies have documented this relationship under natural levels of parasite infestation. We mist-netted 288 adult New Holland honeyeaters (Phylidonyris novaehollandiae) from 12 sites across South Australia to test for a relationship between ticks (Ixodes hirsti) and tail feather asymmetry, and tail feather asymmetry and body condition. Tick prevalence was 15% and intensity varied between 1 and 16 ticks. Tail feather asymmetry did not differ for birds according to their level of tick intensity, or body condition score. At this level of intensity, ticks are unlikely to have had implications for survival over the 4-year study period. Ground feeding and/or nesting birds are potentially exposed to higher levels of tick intensity and would be more suitable for studying the pathogenic effects of I. hirsti.

Nest Predation by Commensal Rodents in Urban Bushland Remnants.

Exotic predators are a major threat to native wildlife in many parts of the world. Developing and implementing effective strategies to mitigate their effects requires robust quantitative data so that management can be evidence-based, yet in many ecosystems this is missing. Birds in particular have been severely impacted by exotic mammalian predators, and a plethora of studies on islands record predation of bird eggs, fledglings and adults by exotic species such as rodents, stoats and cats. By comparison, few studies have examined nest predation around mainland urban centres which often act as dispersal hubs, especially for commensal species such as rodents. Here, we experimentally examine nest predation rates in habitat patches with varying black rat (Rattus rattus) densities in Sydney, Australia and test whether these exotic rats have the effects expected of exotic predators using effect size benchmarks. In the case where black rats have replaced native Rattus spp., we expected that black rats, being more arboreal than native Rattus spp., would be a significant source of predation on birds because they can readily access the arboreal niche where many birds nest. We tested this idea using above-ground artificial nests to represent those of typical small bird species such as the New Holland honeyeater (Phylidonyris novaehollandiae). We found that fewer eggs were depredated by rodents on sites where we removed black rats compared to unmanipulated sites, and that the effect size calculated from the total number of eggs surviving beyond the typical incubation period was similar to that expected for an exotic predator. Our results suggest that, although Australian birds have co-evolved with native Rattus species, in the case where black rats have replaced native Rattus species, exotic black rats appear to pose an additive source of predation on birds in remnant habitats, most likely due to their ability to climb more efficiently than their native counterparts. Management of these commensal rodents may be necessary to retain urban birdlife.

Are blood haemoglobin concentrations a reliable indicator of parasitism and individual condition in New Holland honeyeaters (Phylidonyris novaehollandiae)?

ABSTRACTAcross avian species, total blood haemoglobin concentration (BHC) is the most important determinant of oxygen-carrying capacity, and most accurately reflects the potential for the bird to satisfy its oxygen requirements. This creates a close association between high BHC and high aerobic capacity, and low BHC and states of regenerative or non-regenerative anaemia. As such, total BHC has been suggested to be a reliable indicator of avian health and condition. We mist netted 160 adult and 26 juvenile New Holland honeyeaters (Phylidonyris novaehollandiae) from ten sites across South Australia to assess the relationship between BHC and individual health and condition traits in this species. From each bird we collected samples for blood haemoglobin estimation, inspected for the presence of external parasites (ticks), and measured basic morphometric parameters (mass, tarsus length, and length of bilateral tail feathers). A relationship could not be demonstrated between BHC and tick intensity, body condition or tail feather asymmetry in adult or juvenile birds. While the measurement of BHC may provide a reliable insight into individual health and condition in some avian species, our results highlight the need to validate this relationship within species and populations prior to its use in avian health and condition assessments.

Avian hosts, prevalence and larval life history of the ectoparasitic fly Passeromyia longicornis (Diptera : Muscidae) in south-eastern Tasmania

Blood-sucking fly larvae are widespread parasites of nestling birds, but in many systems we lack knowledge of their basic biology. This study reports the first observation of an endemic Tasmanian fly species, Passeromyia longicornis (Diptera : Muscidae), parasitising the forty-spotted pardalote (Pardalotus quadragintus), another Tasmanian endemic. Because the forty-spotted pardalote is an endangered and declining songbird, P. longicornis is a species of interest to conservation biologists. Its larval form is an obligate, subcutaneous parasite of nestling birds, but before this study, there were just two published records of the species infesting avian hosts, and little known about its ecology or life cycle. This study documented hosts, prevalence, and larval life history of P. longicornis by locating and monitoring nests and ectoparasites of the forest bird community in south-eastern Tasmania. I also reared P. longicornis larvae in captivity to determine the length of the pupal stage in relationship to ambient temperature. Hosts of P. longicornis included forty-spotted pardalotes (87% prevalence across nests), striated pardalotes (Pardalotus striatus) (88% prevalence), and New Holland honeyeaters (Phylidonyris novaehollandiae) (11% prevalence). Both pardalote species were new host records. P. longicornis larvae burrowed under the skin of nestlings where they developed for 4–7 days, feeding on nestling blood. When fully grown, larvae dropped into the surrounding nest material and formed pupae. Length of the pupal stage was 14–21 days, and declined with increasing ambient temperature. Median parasite abundance was 15 larvae in infested forty-spotted pardalote nests and 11 larvae in infested striated pardalote nests. Nestling mortality was frequently associated with ectoparasite presence. This study provides the first survey of P. longicornis hosts, prevalence and life cycle, and shows that this species is likely a major player in the ecology of pardalotes, and possibly other forest bird species in Tasmania.

Does Gray Squirrel (Sciurus carolinensis) Response to Heterospecific Alarm Calls Depend on Familiarity or Acoustic Similarity?

AbstractIn habitats in which multiple species are prey to the same predators, individuals can greatly benefit from recognizing information regarding predators that is provided by other species. Past studies have demonstrated that various mammals respond to familiar heterospecific alarm calls, but whether acoustic similarity to a familiar call can prompt a mammal's recognition of an unfamiliar call has yet to be shown. We presented alarm calls to free‐ranging eastern gray squirrels (Sciurus carolinensis) and recorded behavioral changes in vigilance and antipredatory response. Playbacks included alarm calls of a sympatric bird (American robin, Turdus migratorius), an allopatric bird with a call structure similar to that of the robin (common blackbird, Turdus merula), and an allopatric bird with a distinct call structure (New Holland honeyeater, Phylidonyris novaehollandiae). Squirrels responded significantly more frequently to squirrel alarm calls (positive control) than to robin song (negative control) or honeyeater calls. Squirrel response to robin and blackbird alarm calls was statistically similar to their response to squirrel alarm calls, indicating that squirrels responded to those alarm calls as if they provided information about the presence of predators. However, squirrel response to robin song was not statistically different from response to any of the other avian calls, including the robin and blackbird alarms, suggesting that squirrels neither respond to blackbird alarms as if they clearly signify danger, nor as if they clearly do not signify danger, perhaps reflecting some ambiguity in interpretation of the calls. These results suggest that squirrel responses to alarm calls are generally based on call familiarity, but that acoustic similarity of an unfamiliar allopatric call to a familiar call also can elicit antipredator behavior. The lack of response to honeyeater alarm calls also supports the hypothesis that call recognition by gray squirrels is dependent on familiarity, rather than simply detection of an acoustic feature common to alarm calls across a variety of avian species.

Sugar Preferences of Avian Nectarivores Are Correlated with Intestinal Sucrase Activity

Nectar-feeding birds generally demonstrate preference for hexose solutions at low sugar concentrations, switching to sucrose/no preference at higher concentrations. Species vary in the concentration at which the switch from hexose preference occurs; this could reflect physiological constraints that would also influence nectar selection when foraging. We recorded concentration-dependent sugar type preferences in three opportunistic/generalist Australian nectarivorous species: Dicaeum hirundinaceum, Zosterops lateralis, and Lichenostomus virescens. All three preferred hexoses up to sugar concentrations of 0.25 mol L(-1) and switched to sucrose/no preference for higher concentrations. Using these and literature records, we investigated physiological mechanisms that may explain the concentration dependence of sugar type preferences and compared diet preference data with foraging records. We measured sucrase activity in Z. lateralis and L. virescens as well as three specialized nectarivorous species (Anthochaera carunculata, Phylidonyris novaehollandiae, and Trichoglossus haematodus) for comparison with published concentration-dependent sugar preference data. Sucrase activity varied between these species ([Formula: see text]). The minimum diet concentration at which birds show no sugar preference was significantly correlated with sucrase activity for the 11 species analyzed ([Formula: see text]). Birds with the lowest sucrase activity showed hexose preference at higher diet concentrations, and birds with the greatest sucrase activity showed either no hexose preference or hexose preference on only the most dilute diets. Foraging data compiled from the literature also support the laboratory analyses; for example, T. haematodus (preference for hexose over a wide range of diet concentrations, low sucrase activity) also feed primarily on hexose nectars in the wild. Intestinal sucrase activity is likely to contribute to diet selectivity in nectarivorous bird species.

Eavesdropping on the neighbours: fledglings learn to respond to heterospecific alarm calls

Young birds and mammals suffer from a high risk of predation, and should be under strong selection for early response to cues indicating danger, including the alarm calls of other species. Despite this prediction, there has been little investigation of the development of response by young animals to heterospecific alarm calls, and none on fledgling birds. Previous studies have suggested that learning is important in the recognition of heterospecific alarm calls in adult birds, but when learning occurs is unknown. We examined the responses of fledgling white-browed scrubwrens, Sericornis frontalis, at three ages to playback of the aerial alarm calls of the sympatric superb fairy-wren, Malurus cyaneus, and New Holland honeyeater, Phylidonyris novaehollandiae. Fairy-wren and scrubwren alarm calls are acoustically similar, while honeyeater alarm calls are distinct, but adult scrubwrens respond to both. Recently fledged scrubwrens responded strongly to conspecific alarm calls, but weakly to heterospecific alarm calls. By contrast, only 2 weeks after leaving the nest most fledglings responded to fairy-wren and honeyeater alarm calls similarly to conspecific alarm calls. However, fledglings in territories without honeyeaters ignored honeyeater alarm calls. Three weeks later, fledglings on all territories responded to all heterospecific alarm calls, an identical response to that of their parents. These results demonstrate that fledglings can quickly develop appropriate responses to heterospecific alarm calls, and present strong evidence that response is learned over a microgeographical scale.

Salt intake and regulation in two passerine nectar drinkers: white-bellied sunbirds and New Holland honeyeaters

Avian nectarivores face the dilemma of having to conserve salts while consuming large volumes of a dilute, electrolyte-deficient diet. This study evaluates the responses to salt solutions and the regulation of salt intake in white-bellied sunbirds (Cinnyris talatala) and New Holland honeyeaters (Phylidonyris novaehollandiae). Birds were first offered a choice of four sucrose diets, containing no salt or 25, 50 or 75mM NaCl. The experiment was repeated using five sucrose concentrations (0.075-0.63M) as the base solution. Both species ingested similar amounts of all diets when fed the concentrated base solutions. However, when birds had to increase their intake to obtain enough energy on the dilute sucrose diets, there was a general avoidance of the higher salt concentrations. Through this diet switching, birds maintained constant intakes of both sucrose and sodium; the latter may contribute to absorption of their sugar diets. A second, no-choice experiment was designed to elucidate the renal concentrating abilities of these two nectarivores, by feeding them 0.63M sucrose containing 5-200mM NaCl over a 4-h trial. In both species, cloacal fluid osmolalities increased with diet NaCl concentration, but honeyeaters tended to retain ingested Na(+), while sunbirds excreted it. Comparison of Na(+) and K(+) concentrations in ureteral urine and cloacal fluid showed that K(+), but not Na(+), was reabsorbed in the lower intestine of both species. The kidneys of sunbirds and honeyeaters, like those of hummingbirds, are well suited to diluting urine; however, they also appear to concentrate urine efficiently when necessary.

Learning to listen? Nestling response to heterospecific alarm calls

Many nestling birds go silent in response to parental alarm calls, potentially lowering their risk of being overheard by predators. Parents are not always nearby, however, and so offspring could also benefit if they respond to the alarm calls of other species. Response could be innate, particularly if heterospecific alarm calls are acoustically similar to conspecific alarms, or learned through experience or association with parental behaviour. We investigated the responses of both young (5–6 days old) and older (10–11 days old, close to fledging) nestling white-browed scrubwrens, Sericornis frontalis , to the mobbing alarm calls of three heterospecifics whose nests are vulnerable to similar predators. Brown thornbills, Acanthiza pusilla , produce the most similar alarm calls to scrubwrens, while superb fairy-wren, Malurus cyaneus , and New Holland honeyeater, Phylidonyris novaehollandiae , produce alarm calls that are distinct. Heterospecific mobbing assays demonstrated that nestlings were likely to overhear the alarm calls of all three species. In support of innate response, even young nestlings suppressed calling after hearing both thornbill and parental alarms. However, young nestlings ignored or increased calling to fairy-wren and honeyeater alarms. Older nestlings continued to suppress calling to thornbill and parental alarm calls, but also suppressed calling to honeyeater and fairy-wren alarms, suggesting that they could have learnt to recognize those calls. This study thus demonstrates that nestlings can respond to the alarm calls of other species, and that these responses are likely to be enabled through both innate mechanisms and learning.

Rainfall can explain adaptive phenotypic variation with high gene flow in the New Holland Honeyeater (Phylidonyris novaehollandiae)

Identifying environmentally driven changes in traits that serve an ecological function is essential for predicting evolutionary outcomes of climate change. We examined population genetic structure, sex-specific dispersal patterns, and morphology in relation to rainfall patterns across an island and three peninsulas in South Australia. The study system was the New Holland Honeyeater (Phylidonyris novaehollandiae), a nectarivorous passerine that is a key pollinator species. We predicted that rainfall-related mechanisms would be driving local adaptation of morphological traits, such that in areas of lower rainfall, where nectar is less available, more insectivorous traits – shorter, deeper bills, longer tarsi, and longer wings – would be favored. The study populations differed in phenotype across the Eyre, Yorke, and Fleurieu Peninsulas and Kangaroo Island despite high gene flow (single continuous population) and sex-biased dispersal (males were philopatric and females dispersed). We tested the role of rainfall in shaping the observed phenotypic differences, and found strong support for our predicted relationships: birds in areas of higher rainfall had higher condition indices, as well as longer bill-head length, deeper bills, and shorter tarsi. Bill depth in males in high-rainfall sites showed signals of stabilizing selection, suggesting local adaptation. In addition to these local indications of selection, a global pattern of directional selection toward larger size for bill-head length, bill-nostril length, and wing length was also observed. We suggest this pattern may reflect an adaptive response to the relatively dry conditions that South Australia has experienced over the last decade. We conclude that rainfall has shaped aspects of phenology in P. novaehollandiae, both locally, with different patterns of stabilizing and directional selection, and globally, with evidence of adaptive divergence at a landscape scale.

Do experimental methods affect estimates of pollen digestion by birds?

Pollen protoplasts may supply important nutritional resources for birds; however, they are locked up within the mechanically strong and biochemically complex pollen wall. Previous studies of pollen digestion in birds have yielded highly variable and often contradictory results. We tested whether these differences could reflect the vastly different methodologies that have been used. We used a standard method to investigate digestion of Banksia grandis (Proteaceae) pollen in New Holland honeyeaters (Phylidonyris novaehollandiae). Four types of B. grandis pollen were examined: fresh, frozen for a week, stored in conditions to stimulate pregermination for 24 h, or collected by honeybees. Our data indicate that although pollen treatment may influence digestibility of the pollen grains, these differences do not reach statistical significance because they are dwarfed by a high degree of variability between birds fed the same diet as well as variability in gut transit time (generally more pollen grains were digested over longer transit times). Similar patterns were observed for red wattlebirds (Anthochaera carunculata) fed bee-collected pollen. We believe that feeding behaviour or gut transit time may explain the marked differences between previous studies of pollen digestion by nectarivores, particularly the conflicting results for New Holland honeyeaters.

Added salt helps sunbirds and honeyeaters maintain energy balance on extremely dilute nectar diets

Nectar-feeding birds ingest excess water and risk loss of solutes when they excrete it. Previous work has shown that white-bellied sunbirds (Cinnyris talatala) are unable to maintain energy balance on extremely dilute sucrose diets without salts (e.g. <0.25 mol l(-1)), and that they lose more electrolytes (i.e. Na(+) and K(+)) via cloacal fluid on these diets than on more concentrated diets. Using white-bellied sunbirds and New Holland honeyeaters (Phylidonyris novaehollandiae) we tested the effect of adding electrolytes to a 0.1 mol l(-1) sucrose diet, by including equimolar NaCl and KCl at concentrations from 5 to 40 mmol l(-1) and the individual salts at 20 mmol l(-1). Addition of salts enabled both species to drink significantly more of the 0.1 mol l(-1) sucrose diet than in the absence of salts, and mass loss during the experiment was reduced when salt was included. The larger honeyeaters may be more susceptible to electrolyte depletion than the smaller sunbirds. On 20 mmol l(-1) combined salts, both sunbirds and honeyeaters consumed eight times their body mass in fluid daily. KCl alone had no effect. Birds are thus limited in their consumption of extremely dilute diets by increasing losses of Na(+). This was confirmed by measuring plasma Na(+) levels, which decreased in both species in the absence of dietary Na(+). In addition, sucrose assimilation efficiencies were slightly, but significantly lower when sunbirds were fed salt-free diet, while glucose levels in ureteral urine remained extremely low. It is concluded that Na(+) depletion on very dilute salt-free diets does not affect Na(+)-glucose transport activity in the kidney, but interferes with sugar digestion and/or assimilation in the intestine.

Divergence in New Holland Honeyeaters (Phylidonyris novaehollandiae): evidence from morphology and feeding behavior

Studies of island versus mainland populations have provided insights into processes that shape adaptive divergence (e.g., through selection in allopatric populations) as well as patterns of morphological change. In this study, we compare the morphology and foraging behavior in New Holland Honeyeaters (Phylidonyris novaehollandiae) between Kangaroo Island and the South Australian mainland. We (1) provide descriptive information on morphology traits across populations to test previously described differences; (2) test the prediction that island birds will have a wider foraging niche, different use of foraging substrate, and different foraging techniques, and (3) test for evidence of reduced prey diversity on islands as evidenced by longer foraging time. The results provide support for the predictions. Island birds were larger than the mainland birds in tarsus (2.5%) and bill length (3.7%), had a wider foraging niche (mostly due to greater insect consumption), and foraged more from the bark and air (sallying). Consistent with other studies of island biogeography, at least one common mainland predator and several bird species are absent from Kangaroo Island, which may facilitate niche expansion in the island birds. A wider niche could also be favored by lower prey diversity on islands (a depauperate community): in this study, island birds had longer foraging times than mainland birds, which may be evidence for lower total prey density.