Trophic cascades in bell miner‐associated dieback forests: Quantifying relationships between leaf quality,psyllids and Psyllaephagus parasitoids

Coteries of the meliphagid bird Manorina melanophrys are associated with a form of eucalypt defoliation and recovery called bell miner‐associated dieback (BMAD). Through their defence of cooperative colony boundaries against other insectivorous birds, bell miners may foster greater abundances of lerp‐forming psyllids (Hemiptera: Aphalaridae), some of which reduce the lifespan of leaves. Trophic cascades in BMAD forests need to be understood to have a complete picture of regulatory processes. We studied relationships between leaf quality, psyllid and Psyllaephagus parasitoid/hyperparasitoid abundances within the Gondwana Rainforest World Heritage Area, NSW, Australia; our focal tree species were Eucalyptus propinqua and E. biturbinata. Eucalyptus biturbinata had tougher leaves than E. propinqua; leaf toughness of both species varied with site and tree. We found a statistically significant, negative relationship between toughness (surrogate for leaf age) and foliar nitrogen content; younger leaves had higher nitrogen contents. Both bell miner abundance and foliar nitrogen were positively correlated with psyllid abundance. The abundance of Glycaspis species (the psyllid that produces lerps with the highest sugar content) was more closely correlated with foliar nitrogen content than was the abundance of all five psyllid genera combined. We identified 14 Psyllaephagus spp./morphospecies, comprising 11 primary parasitoids and three hyperparasitoids. The abundance of all Psyllaephagus combined was positively correlated with the abundance of lerps. However, psyllid parasitism was not correlated with the abundance of lerps. The abundance of the three hyperparasitoids was positively correlated with the abundance of Psyllaephagus hosts. The availability of epicormic foliage (young, morphologically juvenile leaves produced following defoliation) is likely to alter the nutritional ecology underpinning the diversity and abundance of psyllid populations. Higher quality epicormic foliage should favour populations of Glycaspis species (by enhancing nymphal survival) creating lerp hotspots that induce residency by opportunistic bell miners. The positive contribution of induced amelioration, interacting with feedbacks from parasitoids and hyperparasitoids, to BMAD requires longitudinal investigation.     

Fine‐scale impacts on avian biodiversity due to a despotic species,the bell miner (Manorina melanophrys)

Bell miners (Manorina melanophrys; Meliphagidae) are a highly social and very aggressive honeyeater. They are despotic and cooperate in the defence of their territories against other bird species, leading to the almost complete exclusion of other avifauna from miner‐occupied regions. This study aimed to resolve some of the fine‐scale effects of bell miner aggression on avian diversity both within and adjacent to colonies to determine the true impact of a colony on local avifaunal abundance. Three areas, distributed throughout the range of the bell miner, were surveyed across both non‐breeding and breeding seasons to assess the temporal and spatial impacts of bell miner aggression on other bird species. Bell miner colonies were found to occupy very clearly defined areas and had the expected negative impact on avian diversity within their colony. The effects of bell miner colony presence on abundance and richness of avian species were found to cease at the colony boundary, with both recovering to normal levels immediately outside the bell miner colony. Whether bell miners were breeding or not, and irrespective of the amount of vegetation coverage, bell miner colonies were found to have relatively marginal impacts on avian richness and abundance. No impact of colony presence/absence was found on the richness or abundance of the avian species that dwell in the undergrowth, with some evidence that these species were actually more common at the colony edge. Our results demonstrate that the influence of bell miner colony presence upon avian biodiversity is restricted to the confines of the colony and does not radiate outwards into the surrounding habitat. Colony presence influences, therefore, have implications when considering the impact of bell miner behaviour on the diversity of insectivorous birds and processes, most notably the propagation of Bell Miner Associated Dieback.     

Impacts of grazing,selective logging and hyper‐aggressors on diurnal bird fauna in intact forest landscapes of the Brigalow Belt,Queensland

The impact of forest management on diurnal bird assemblages and abundance was investigated in contiguous tracts of eucalypt forest in the Brigalow Belt Bioregion, south central Queensland. Sites were located across three levels of livestock grazing intensity and three levels of selective logging intensity within the most extensive habitat type, Corymbia citriodora‐dominant forest. We recorded a high rate of incidence and large numbers of the hyper‐aggressive noisy miner Manorina melanocephala (Passeriformes: Meliphagidae) at the majority of our survey sites, a phenomenon rarely reported in non‐cleared landscapes. As shown by numerous studies in fragmented landscapes, the distribution of this species in our study had a substantial negative effect upon the distribution of small passerine species. Noisy miners exerted the strongest influence upon small passerine abundance, and masked any forest management effects. However, key habitat features important for small passerines were identified, including a relatively high density of large trees and stems in the midstorey. Selective logging appeared to exert a minimal effect upon noisy miner abundance, whereas grazing intensity had a profound, positive influence. Noisy miners were most abundant in intensively grazed forest with minimal midstorey and a low volume of coarse woody debris. Higher road density in the forest landscape also corresponded with increased numbers of noisy miners. Reduction in grazing pressure in Brigalow Belt forests has the potential to benefit small passerine assemblages across large areas through moderating noisy miner abundance. The strong relationship between noisy miners and small passerines suggests that noisy miner abundance could act as an easily measured indicator of forest condition, potentially contributing to monitoring of forest management outcomes.     

A particular case and a general pattern: hyperaggressive behaviour by one species may mediate avifaunal decreases in fragmented Australian forests

We quantitatively assessed edge effects associated with elevated abundance of a hyper aggressive bird species, the noisy miner Manorina melanocephala, in fragmented eucalypt forest adjoining developed land. Long‐term data from Toohey Forest, subtropical Australia, show that noisy miner colonies intensively occupy a zone of 20 m from the forest edge, with frequent use occurring up to 100 m from the edge, but little beyond 200 m. Within noisy miner colonies, the abundance and species richness of other birds were both about half those recorded at nearby transects which were outside the colonies' main activity area. Bird species smaller than noisy miners, which are also those with similar diets, were collectively 20–25 times more abundant, and their species richness tenfold greater, outside miner colonies than within them, whereas larger species, which have less dietary overlap, did not differ. Exclusion of small insectivorous birds has been hypothesised to cause elevated insect herbivore density, but we found no difference between tree crown defoliation or dieback rates within versus outside miner colonies. Aggression by noisy miners can be viewed as a mechanism of interspecific competition, since miners have a relatively large body size for their diet and are hence able to exclude virtually all potential competitors at relatively little cost. We examine evidence indicating that reduced bird diversity in eucalypt forest fragments of eastern Australia is often simply the effect of noisy miner occupancy of edges, acting directly on the densities of other species through their aggressive behaviour. With an edge effect 200 m deep, a remnant 10 ha in size is likely to become entirely occupied by noisy miners, and this is a size threshold that has been commonly reported in association with area‐standardised avian diversity reductions. Convergent patterns of species loss from small forest fragments in different continents are the result of different underlying ecological processes.     

The portability of foodweb dynamics: reassembling an Australian eucalypt–psyllid–bird association within California

Aims To evaluate the role of native predators (birds) within an Australian foodweb (lerp psyllids and eucalyptus trees) reassembled in California. Location Eucalyptus groves within Santa Cruz, California. Methods We compared bird diversity and abundance between a eucalyptus grove infested with lerp psyllids and a grove that was uninfested, using point counts. We documented shifts in the foraging behaviour of birds between the groves using structured behavioural observations. Additionally, we judged the effect of bird foraging on lerp psyllid abundance using exclosure experiments. Results We found a greater richness and abundance of Californian birds within a psyllid infested eucalyptus grove compared to a matched non‐infested grove, and that Californian birds modify their foraging behaviour within the infested grove in order to concentrate on ingesting psyllids. This suggests that Californian birds could provide indirect top‐down benefits to eucalyptus trees similar to those observed in Australia. However, using bird exclosure experiments, we found no evidence of top‐down control of lerp psyllids by Californian birds. Main conclusions We suggest that physiological and foraging differences between Californian and Australian pysllid‐eating birds account for the failure to observe top‐down control of psyllid populations in California. The increasing rate of non‐indigenous species invasions has produced local biotas that are almost entirely composed of non‐indigenous species. This example illustrates the complex nature of cosmopolitan native‐exotic food webs, and the ecological insights obtainable through their study.     

The role of psyllids (Hemiptera: Psyllidae) and bell miners (Manorina melanophrys) in canopy dieback of Sydney blue gum (Eucalyptus saligna Sm.)

Abstract The presence of the aggressive, colony-forming honeyeater, Manorina melanophrys (bell miner), in the canopies of unhealthy eucalypts has been well reported. There is, however, some debate as to the actual mechanisms producing these unhealthy trees. To investigate further some of the processes that may be contributing to this form of canopy dieback, two field trials were carried out in Olney State Forest, near Wyong, New South Wales. The study site contained Eucalyptus saligna (Sydney blue gum) with canopy dieback and was occupied by a large colony of bell miners. Close examination of the foliage revealed a large and diverse suite of phytophagous insects, including at least 16 species of psyllid (Hemiptera: Psyllidae). In the first trial, the use of bird exclusion cages over selected branches significantly improved leaf survival compared to leaves exposed to a relatively high density of bell miners. It is proposed that colonization by bell miners may interfere with the efficacy of both other insectivorous birds (through aggressive interspecific territoriality) and the invertebrate predators and parasitoids. Interference with such regulatory factors may enable some phytophagous insect populations to rise to sustained damaging levels. In the second trial, an insecticide application combined with reduced competition from the dense understorey and neighbouring trees was required to significantly improve trunk diameter and crown condition scores. After 12 months, neither treatment, by itself, significantly improved both growth measures. Possibly both treatments were required because the E. saligna trees were suffering from another source of stress (e.g. drought) in addition to the relatively high level of insect attack.     

Sustained and delayed noisy miner suppression at an avian hotspot

To mitigate the impact of noisy miners Manorina melanocephala on Australia’s woodland birds, there is a need to identify locations where noisy miner suppression can be affordable, sustainable and facilitate woodland bird recovery. In 2017, we suppressed noisy miners from the Goulburn River, NSW for at least three months. During this period, six pairs of critically endangered regent honeyeaters nested in the treatment area. In 2018, we continued monitoring the original noisy miner treatment area, which was expanded to include our 2017 control area, and established a new control area downstream. In 2019, the removal effort was again expanded to include the 2018 control area. In the 2017 treatment area, noisy miners remained suppressed up to 27 months post‐removal. Their numbers here were lower 1 year after the initial cull than in the week after it. In the 2018 and 2019 treatment areas, noisy miner abundance was significantly lower after respective culls than at all pre‐removal periods. In 2018, around 20 vulnerable painted honeyeaters occupied the 2018 treatment area. In 2019, two regent honeyeater pairs nested in and at least 40 painted honeyeaters occupied the treatment area. Songbird abundance increased within seasons and also up to a year following noisy miner removal, and plateaued thereafter. We show how, in strategic locations, a week of noisy miner suppression each spring can sequentially create ever‐larger landscapes where noisy miner impacts on threatened woodland birds are minimal.     

Short‐term response of a declining woodland bird assemblage to the removal of a despotic competitor

Interspecific aggression by the noisy miner (Manorina melanocephala), a highly despotic species, is homogenizing woodland avifaunas across eastern Australia. Although a native species, the noisy miner's aggressive exclusion of small birds is a Key Threatening Process under national law. Large‐scale removal of noisy miners has been proposed as a management response to this threat following increases in miner presence due to anthropogenic land use practices. We tested this proposal by experimentally removing noisy miners from eucalypt woodland remnants (16–49 ha), assigned randomly as control (n = 12) or treatment (miner removal) sites (n = 12). Standardized bird surveys were conducted before and after removal, and generalized linear mixed models were used to investigate the effect of miner removal on bird assemblage metrics. Despite removing 3552 noisy miners in three sessions of systematic shooting, densities of noisy miners remained similarly high in treatment and control sites, even just 14 days after their removal. However, there was evidence of an increase in richness and abundance of small birds in treatment sites compared to controls—an effect we only expected to see if noisy miner densities were drastically reduced. We suggest that miner removal may have reduced the ability of the recolonizing miners to aggressively exclude small birds, even without substantially reducing miner densities, due to the breakdown of social structures that are central to the species' despotic behaviour. However, this effect on small birds is unlikely to persist in the long term. Synthesis and applications: Despite evidence from other studies that direct removal of noisy miners can result in rapid and sustained conservation benefit for bird communities at small scales, our findings cast doubt on the potential to scale‐up this management approach. The circumstances under which direct control of noisy miners can be achieved remain unresolved.     

Development of Glycaspis brimblecombei Moore (Hemiptera: Aphalaridae) on Eucalyptus camaldulensis Dehnh. and Eucalyptus dunnii Maiden

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Boom and bust: rapid feedback responses between insect outbreak dynamics and canopy leaf area impacted by rainfall and CO2

Frequency and severity of insect outbreaks in forest ecosystems are predicted to increase with climate change. How this will impact canopy leaf area in future climates is rarely tested. Here, we document function of insect outbreaks that fortuitously and rapidly occurred in an ecosystem under free‐air CO₂ enrichment. Over the first 2 years of CO₂ fumigation of a naturally established mature Eucalyptus woodland, we continuously assessed population responses of three sap‐feeding insect species of the psyllid genera Cardiaspina, Glycaspis and Spondyliaspis for up to ten consecutive generations. Concurrently, we quantified changes in the canopy leaf area index (LAI). Large and rapid shifts in psyllid community composition were recorded between species with either flush (Glycaspis) or senescence‐inducing (Cardiaspina, Spondyliaspis) feeding strategies. Within the second year, two psyllid species experienced significant and rapid population build‐up resulting in two consecutive outbreaks: first, rainfall stimulated Eucalyptus leaf production increasing LAI, which supported population growth of flush‐feeding Glycaspis without impacting LAI. Glycaspis numbers then crashed and were followed by the outbreak of senescence‐feeding Cardiaspina fiscella that led to significant defoliation and reduction in LAI. For all three psyllid species, the abundance of lerps, protective coverings excreted by the sessile nymphs, decreased at e[CO₂]. Higher lerp weight at e[CO₂] for Glycaspis but not the other psyllid species provided evidence for compensatory feeding by the flush feeder but not the two senescence feeders. Our study demonstrates that rainfall drives leaf phenology, facilitating the rapid boom‐and‐bust succession of psyllid species, eventually leading to significant defoliation due to the second but not the first outbreaking psyllid species. In contrast, e[CO₂] may impact psyllid abundance and feeding behaviour, with psyllid species‐specific outcomes for defoliation severity, nutrient transfer and trophic cascades. Psyllid populations feeding on Eucalyptus experience rapid boom‐and‐bust cycles depending on availability of suitable foliage driven by rainfall patterns and leaf phenology.     

Chronic stress in superb fairy‐wrens occupying remnant woodlands: Are noisy miners to blame?

Interactions between competing species may be intensified when they are restricted to small patches of remnant habitat, potentially increasing physiological stress in individuals. The effects of interspecific competition on stress in wildlife remain largely unexplored. In Australia, remnant woodlands are often dominated by aggressive honeyeaters, especially the noisy miner (Manorina melanocephala). Harassment of smaller birds by miners may result in their exclusion from suitable woodland habitat. We tested whether the presence of noisy miners is also associated with elevated stress in a model species of small passerine bird, the superb fairy‐wren (Malurus cyaneus). We sampled wrens from six sites, three remnant woodlands with noisy miners and three larger fragments of reserved habitat without noisy miners. Differential white blood cell counts were used to infer levels of chronic stress. We also assessed variation in body condition and the prevalence of blood parasites (Haemoproteus spp.) to test for associations between stress and parasitemia. The mean heterophil‐to‐lymphocyte (H:L) ratio was 1.8 × higher among superb fairy‐wrens living in miner‐dominated woodlands, suggesting higher levels of chronic stress. Individuals with higher stress appeared to be in poorer condition, as indicated by fat scores and residual body mass. Prevalence of blood parasites was generally high and was highest in reserved habitat (59%) where miners were absent. Birds with blood parasites living in these habitats had higher H:L ratios but the intensity of infection and H:L ratio was inversely related. Our results suggest that birds persisting in the presence of noisy miners might experience chronic stress, but further study is necessary to separate the relative importance of noisy miner aggression from other potential stressors in small patches of degraded woodland. Stress induced by interspecific aggression should be considered in future studies of wildlife living in remnant vegetation.     

Habitat preferences of the noisy miner (Manorina melanocephala) – a propensity for prime real estate?

This study investigated habitat characteristics that have been postulated to influence the occurrence of noisy miners (Manorina melanocephala Family Meliphagidae). It builds on an earlier study that identified corners along remnant edges as important predictors of the presence of noisy miners in large blocks of remnant vegetation (>300 ha). Six habitat characteristics were recorded at 39 corner sites within the box‐ironbark region of Victoria. We failed to detect any significant effect of the density of understorey vegetation on the likelihood of noisy miners occupying a site, but this might have been an artefact of prolonged drought conditions. The most powerful predictors of the presence of noisy miners at remnant corners were found to be soil type and the proportion of canopy trees at a site that were yellow gum (Eucalyptus leucoxylon), with noisy miners being associated with deeper, more fertile soils and higher proportions of yellow gums present. As yellow gum is a prolific and reliable nectar producer, the inherent productivity of a site might be more important in determining the attractiveness of a site to noisy miners than structural attributes like the presence or absence of an understorey. Noisy miners are sedentary colonial birds that occupy year‐round territories, often at high densities. Sites capable of supporting such high density occupation year‐round might be limited to the most productive sites within the landscape. This productivity hypothesis has potentially profound implications for other woodland avifauna, as noisy miners might be excluding other woodland birds from some of the most fertile components of the landscape; components that are already rare in the box‐ironbark region due preferential clearing for agriculture at the time of settlement.     

Group Mobbing Behaviour and Nest Defence in a Cooperatively Breeding Australian Bird

Cooperatively breeding noisy miners (Manorina melanocephala) are well known in Australia for their persistent and very vocal group mobbing of heterospecifics. Here I investigated the nature of this extraordinary behaviour, in particular its role in nest defence, in a colour banded population of noisy miners in south‐east Queensland, Australia. I focused on two questions. First, did the intensity of mobbing vary according to factors such as the threat to the nest, or the ‘value’ of a clutch? Secondly, what role did group mobbing play in the success of a nest? To answer these questions, I experimentally manipulated the nest defence behaviour by placing one of three stuffed models near active noisy miner nests. The response of noisy miners to intruders was not indiscriminate. However, I found that the number of birds that mobbed a model did not simply reflect the potential threat posed. The response of noisy miners to raptors and other potential nest predators may have reflected their rarity as well as the threat posed. The number of mobbers did not vary with the age or size of a brood. In this study, the fate of nests was independent of the number of mobbers or visitors at nests. Finally, up to 80% of mobbers were never seen to make any other type of contribution to a nest, and many could not be related to the brood that they were ‘defending’. Hence, for some noisy miner ‘helpers’ the benefits that they accrued were probably not wholly dependent on the survival of the broods. I suggest that, in this gregarious species, mobbing behaviour at the nest may be a display of social status or individual quality. This hypothesis warrants further investigation.     

Yellow‐throated miners Manorina flavigula homogenize bird communities across intact and fragmented landscapes

In Australia, the role of noisy miners Manorina melanocephala in biotic homogenization of the avifauna has been well established in modified landscapes, and is listed as a threatening process under national conservation legislation. However, less is known about the effect of the congeneric and more widely distributed yellow‐throated miner, M. flavigula. In this paper we investigate the relative roles of habitat loss and increased dominance by the yellow‐throated miner in avian homogenization and species functional group decline. We examined bird community data collected from 368 woodland sites across three bioregions. For each site there was a local and a landscape scale measure of remnant vegetation cover. We used both multivariate and regression analysis to test the relative influence of yellow‐throated miner abundance and vegetation on bird community composition. There was clear compositional change and homogenization of the avifauna where yellow‐throated miners were present and vegetation cover was low. The abundance of 40 bird species was predicted by combinations of vegetation cover or yellow‐throated miner abundance, and 31 of these regressions included the term yellow‐throated miner. Of these, there was a negative relationship with 23 species, and 19 of these were insectivores or nectarivores. We postulate that the combination of clearing and yellow‐throated miner abundance can interact to disrupt the ecological function of woodlands, by the depletion of insect‐ and nectar‐feeding species and the disturbance to mixed feeding flocks. We propose future research objectives that include a continental‐scale analysis of the determinants of yellow‐throated miner overabundance, the numerical and geographical thresholds of their potential impacts, and the ecological consequences on both avifauna and the woodlands they inhabit.     

Do miners (Manorina spp.) affect bird assemblages in continuous savanna woodlands in north‐eastern Australia?

Overabundant native species can have a significant cascading effect on other components of wildlife, and those that deplete other species, often promoted by anthropogenic change to vegetation cover and habitat, are called reverse keystone species. Birds in the genus Manorina are widely reported as being such species, and in highly disturbed or fragmented environments, and some intact environments, noisy miners Manorina melanocephala can have a strong negative effect on small passerine species via hyper‐aggressive mobbing. The tropical savannas of northern Australia consist of largely unmodified woodlands, and two species of Manorina occur naturally in this region: the noisy miner and the yellow‐throated miner Manorina flavigula. Therefore, what effect do these species have on bird assemblage in predominantly continuous habitats, relative to other typical determinants of avifauna assemblage such as vegetation structure? We used data collected from bird surveys at 511 sites across northern Queensland (179 noisy miner M. melanocephala sites, 332 yellow‐throated miner M. flavigula sites) between 1998 and 2010. We examined the variation in bird composition at each site due to increasing abundance of Manorina spp. using uni‐ and multivariate techniques. We found total bird richness was significantly lower in sites where noisy and yellow‐throated miner abundances were highest, and passerine species seemed most affected. For species, 45 species varied significantly in abundance with increasing miner numbers, and the overall effect of yellow‐throated miners on other birds seemed more pronounced. However, vegetation structure was generally an equal or more important predictor of avifauna richness and abundance. We conclude that despite the superficially intact nature of northern Australian woodlands, pastoral intensification or poor land management might create disturbances that facilitate increases in the abundance of Manorina, causing localized overabundance and a compounding negative effect on other native bird species.     

Effect of host deprivation on the foraging behavior of the Asian citrus psyllid parasitoid Tamarixia radiata: observations from the laboratory and the field

The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae), is a significant citrus pest and the parasitoid Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae) has been released in various citrus‐producing areas in classical biological control programs targeting D. citri. We investigated the effect of host deprivation on the foraging behavior and patch utilization by T. radiata. In the laboratory, females deprived of hosts for 3 days tended to leave patches of 12 nymphs without parasitizing hosts during the ca. 30 min they spent in the patch before leaving. Moreover, half of these females failed to host feed, and those that did host feed, on average, needed more than 15 min to complete feeding. Conversely, non‐host‐deprived females parasitized on average three nymphs before leaving patches without host feeding during the ca. 39 min they spent in the patch. These laboratory observations were compared to mass‐reared female T. radiata that were released onto colonies of D. citri nymphs infesting citrus in the field. Release vials were provisioned with honey and these females had no opportunities to host feed over the 1‐ to 2‐day containment period prior to release. When introduced onto D. citri colonies, 68% T. radiata females abandoned D. citri patches prior to probing hosts, in part, because Argentine ants, Linepithema humile (Mayr) (Hymenoptera: Formicidae), tending colonies disturbed searching parasitoids. These results from laboratory and field studies are discussed in the context of classical biological control, with the aim of understanding how to manipulate host availability and ant activity so establishment rates and impact of T. radiata can be improved.     

Sampling and interpretation of psyllid nymph counts in potatoes

Development of effective management practices for insect pests relies heavily on sampling methods to accurately detect and quantify emerging populations. Herein we describe a novel method to extract and count nymphs of potato psyllid, Bactericera cockerelli (?ulc) (Hemiptera: Triozidae), from leaves in commercial fields of potato, Solanum tuberosum L. (Solanaceae). The proposed sampling method (referred to as the leaf washing method, LWM) consists of: (1) immersing samples of infested leaves in cold water to remove dust and sand; (2) immersing leaves in >85 °C water for 5 s; (3) extracting psyllid nymphs from heated water by passing it through a sampling unit composed of a funnel with a fine mesh‐organza fabric inside and attached to a vacuum pump; and (4) removing organza fabric and count psyllid nymphs under a stereoscope. With five sampling units operating simultaneously, one person is able to process and count immatures from about 400 leaf samples in two normal work days. As examples of applications, the LWM was used to characterize: (1) the vertical distribution of potato psyllid nymphs in the potato canopy; and (2) the spatio‐temporal distribution of potato psyllid nymphs in three potato varieties. Using LWM, we showed that psyllid nymphs were most predominant in the middle portion of the canopy and that spatio‐temporal distributions of nymphs varied among potato varieties.     

Facultative control of offspring sex in the cooperatively breeding bell miner, Manorina melanophrys

The ability to alter primary sex ratios has the potential toincrease a breeding individual's fitness. This is certainlytrue in those cooperative breeders where one sex is both philopatricand helps raise future offspring of its parents. We examinedthe primary sex ratio variation in the cooperatively breedingbell miner (Manorina melanophrys) in southeastern Australiaover six breeding seasons. Male offspring are the philopatricand helping sex in this system and can increase the reproductiveoutput of their parents. Bell miners aggressively defend theirterritory from all interspecific competitors and by doing soallow food resources to dramatically increase. The increasein phytophagous Psyllidae insects (which secrete a carbohydrate-richcoating that constitutes the major component of bell miner diet)leads to a decrease in tree health, often culminating in deathof the tree. Bell miners then move as groups to new areas withlow psyllid abundance, and the cycle repeats. Using this predictabletemporal variation in food availability, we aimed to determinewhether female breeders adjusted the sex ratio of broods toproduce more of the philopatric sex when food resources werehigh and more of the dispersing sex when food resources werelow. Our results provide clear evidence for such facultativecontrol of sex ratio by female bell miners. Newly founded coloniesare characterized by low food availability and a female-biasedprimary sex ratio, whereas colonies more than 1 year old havean increased food availability and a male-biased primary sexratio. We suggest treating forces associated with resource enhancementand competition as opposing sides of a single general principleand suggest that it is necessary to view both the costs andbenefits of philopatric individuals within a variable environment.     

The effects of conspecific and heterospecific interactions on foraging and oviposition behaviours of two parasitoids of Diaphorina citri

Direct observation was used to construct activity budgets for Diaphorencyrtus aligarhensis (Shafee, Alam & Agarwal) (Hymenoptera: Encyrtidae) and Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae), foraging for nymphs of Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae). Two populations of D. aligarhensis were used in experiments: a biparental population from Pakistan and a uniparental population from Taiwan. Parasitoids were provisioned with 20 D. citri nymphs, second through fifth instar; female parasitoids from all three populations foraged alone, in conspecific pairs, or in heterospecific pairs. Walking occupied approximately 40% of the activity budget of D. aligarhensis and ovipositor probing occupied 10–20%. In contrast, probing and walking each accounted for approximately 30% of the activity budget of T. radiata. The presence of conspecific females affected the activity budget of D. aligarhensis females, such that the focal female was observed walking less often and probing more frequently. Heterospecific females did not affect D. aligarhensis activity. These results suggest mutual interference may occur between D. aligarhensis females foraging on the same host patch. The behaviour of T. radiata was not significantly affected by the presence of conspecific or heterospecific competitors. These results are discussed with respect to the release of multiple species of natural enemies in the classical biological control programme targeting D. citri in California.