Biocontrol without borders: the unintended spread of introduced weed biological control agents

An underlying assumption of classical biological control implies that intentionally introduced natural enemies will remain within the boundaries that delineate the program’s area of implementation. A weed biological control program targeting Melaleuca quinquenervia in Florida, USA has resulted in the release and establishment of Oxyops vitiosa and Boreioglycaspis melaleucae. An international survey of M. quinquenervia populations in 13 other states or countries where the insects have not been intentionally introduced was initiated to monitor the long range dispersal of O. vitiosa and B. melaleucae beyond the herbivores’ intended geographic range (Florida). Surveys in 2006 resulted in the discovery of B. melaleucae within the canopies of several M. quinquenervia trees near San Juan, Puerto Rico. In 2007, O. vitiosa was observed on the island of New Providence in the Bahamas but neither herbivore was detected on nearby Grand Bahama or Andros islands. In 2009, B. melaleucae was observed attacking M. quinquenervia trees in Los Angeles, California (USA). The herbivores have not been detected on other surveyed M. quinquenervia populations in Cuba, Jamaica, Texas (USA), Costa Rica, Brazil, Hawaii (USA) or South Africa. There is no evidence to suggest that herbivore colonization of New Providence, Puerto Rico, or California was influenced by linear distance between Florida and the recipient M. quinquenervia stand. While the dispersal pathway(s) remains unknown, biological control agents were detected from 200 to >3500 km from their original release location (Florida) and at locations that have strong links via tourism and trade as indicated by the number of airline flights connecting south Florida with colonized tree populations. Implications of this unintended spread are discussed in relation to permeability of biogeographical barriers and risk assessment of biological control agents.     

Biological control increases the susceptibility of melaleuca quinquenervia to fire

A field experiment evaluating herbivory from biological control agents on Melaleuca quinquenervia saplings experienced a wildfire, which resulted in post-fire tree mortality of 64.6% and 28.1% in the unrestricted and restricted herbivory treatments, respectively. The fire did not negatively impact populations of the biological control agent Oxyops vitiosa.     

Antipredatory Activity of the Weevil Oxyops vitiosa: A Biological Control Agent of Melaleuca quinquenervia

The larvae of the leaf-feeding weevil Oxyops vitiosa, a biological control agent of Melaleuca quinquenervia, are covered with a viscous orange coating that is thought to protect against generalist predators. This coating is gradually lost as the larvae drop to the ground and pupate in subterranean pupal cells. To test the antipredator activity of this species, four immature life stages (early instars, late instars, prepupae, pupae) were exposed to a common generalist predator, the red imported fire ant Solenopsis invicta. Choice tests were conducted by placing an O. vitiosa individual and a control larva of the weevil Neochetina eichhorniae into an arena containing a S. invicta colony and observing subsequent ant behaviors. S. invicta workers contacted O. vitiosa early instars, late instars, and prepupae less frequently than control N. eichhorniae larvae, and upon contact S. invicta was less likely to behave aggressively toward these O. vitiosa life stages than toward N. eichhorniae larvae. However, S. invicta contacted, attacked, and consumed naked (nonencased) O. vitiosa pupae and N. eichhorniae larvae with equal frequency. Encased O. vitiosa pupae buried in sand were not attacked compared to susceptible encased pupae on the sand surface. By shifting from a chemical defense during the larval stages to a physical defense during the pupal stage, O. vitiosa reduces the risk of attack by this generalist predator.     

Herbivory alters resource allocation and compensation in the invasive tree Melaleuca quinquenervia

Abstract. 1. Plants may compensate for the effects of herbivory, especially under favourable growing conditions, limited competition, and minimal top‐down regulation. These conditions characterise many disturbed wetlands dominated by introduced plants, implying that exotic, invasive weeds in these systems should exhibit strong compensatory responses. 2. The Australian native Melaleuca quinquenervia is highly invasive in the Florida Everglades, U.S.A., where it experiences limited competition or herbivory from native species, making it a likely candidate for compensation. The introduced biological control agent Oxyops vitiosa feeds exclusively on the seasonal flushes of developing foliage at branch apices, which represents ≈15% of the total foliar biomass. 3. The hypothesis that M. quinquenervia compensates for folivory by O. vitiosa was tested in a series of field‐based experiments. Trees experiencing folivory over four consecutive years maintained similar levels of foliar biomass after attack yet possessed twice the number of leaf‐bearing terminal stems as undamaged trees. The biomass of these stems was similar among attacked and unattacked trees, indicating that herbivore‐damaged trees produce greater quantities of smaller terminal branches. However, undamaged trees were 36 times more likely to reproduce than herbivore‐damaged trees. 4. In a separate herbivore exclusion study, a single bout of herbivory on previously undamaged M. quinquenervia trees caused an 80% reduction in reproductive structures the following season. Herbivore‐damaged trees also possessed 54% fewer fruits than undamaged trees. An increase in the herbivory frequency (two bouts per year) or magnitude (100% simulated herbivory) did not result in a further reduction in fitness. 5. It has been concluded that M. quinquenervia partially compensates for herbivory by producing new stems and replacing foliage, but this compensation results in a substantial reduction in reproduction.     

Modeling the Influence of Resource Availability on Population Densities of Oxyops vitiosa (Coleoptera: Curculionidae), a Biological Control Agent of the Invasive Tree Melaleuca quinquenervia

The Australian snout beetle, Oxyops vitiosa, was introduced to south Florida, USA, as a biological control agent of the invasive tree Melaleuca quinquenervia during the spring of 1997. As part of ongoing post-release evaluations of this weevil, we sought to quantify the population density that can be maintained by O. vitiosa larvae when fully exploiting the available melaleuca foliage. Seasonal population densities were modeled as a function of tree size distribution and density, plant phenology, quantity of acceptable foliage (suitable for larval development) and larval consumption values. Larval densities varied dramatically due to fluctuating resource availability and ranged from 830 946 larvae/ha in July–August to >4.5 million larvae/ha in January–February. The expected population density for a generalized (mixed sized) tree distribution was estimated to be 13.9 million larvae/ha. Larval densities increased to 18.8 million larvae/ha per year when large trees (>20 cm diameter at breast height) represented the dominant size class, whereas habitats dominated by medium (10–19 cm) and small (1.3–9 cm) trees were predicted to have larval densities of 11.3 and 6.3 million/year, respectively. Validation data obtained from smaller, more accessible plants suggested that the model overestimated realized yearly larval densities by 9% or 15 416 larvae/ha per year.     

Nonselective oviposition by a fastidious insect: the laboratory host range of the melaleuca gall midge Lophodiplosis trifida (Diptera: Cecidomyiidae)

Invasion by the Australian paperbark, Melaleuca quinquenervia, has degraded large areas of south Florida wetlands. Restoration of these wetlands requires the removal of expansive monocultures of this large tree while simultaneously curtailing its spread. Management strategies developed by federal and state agencies include biological control to halt the spread of this species and to prevent reinfestation of cleared areas. This requires biological agents able to reduce flowering, seed production, and growth while increasing mortality of seedlings and saplings. Two of the three introduced agents (Oxyops vitiosa Pascoe and Boreioglycaspis melaleucae Moore) partially meet these needs but outcomes are not spatially or temporally consistent. Thus, additional agents are needed. The bud-gall fly Fergusonina turneri Taylor, with its mutualistic nematode Fergusobia quinquenerviae Davies and Giblin-Davis, is actively being released but has not established. A fourth promising agent, the gall midge Lophodiplosis trifida Gagné, manifested an extremely narrow host range during laboratory testing. Oviposition was indiscriminant in caged environments. Small, incipient, unilocular galls were initiated on Melaleuca viminalis, but larval development ensued only on M. quinquenervia. The unilocular galls on M. viminalis did not grow and produced no adult flies. As a result, M. viminalis test plants suffered only minor cosmetic damage. Observations from both Australia and Florida attest to the ability of this midge to impede M. quinquenervia growth and kill small plants. Thus, L. trifida is safe to release and will likely contribute to management objectives for control of this pernicious wetland invader.     

Resource regulation of an invasive tree by a classical biological control agent

The invasive tree Melaleuca quinquenervia experienced substantial declines in growth and reproduction in response to chronic herbivory by the defoliating weevil Oxyops vitiosa. Plants subjected to unrestricted defoliation replaced leaves that were more suitable for feeding by the next generation, a process envisioned by the Resource Regulation Hypothesis which posits that attack by one generation increases the amount of the preferred host resources for the next, resulting in a positive feedback loop for the herbivore. The production of juvenile replacement leaves stimulated additional bouts of oviposition and feeding by O. vitiosa, which ultimately produced positive effects for the herbivore with negative consequences for the plant. The addition of water resources to the plant prolonged the positive feedback loop such that more than twice as many insects were produced on irrigated versus non-irrigated trees. In a more simple, reassembled food web on M. quinquenervia, the lack of biotic constraints like parasitoids may have prevented the earlier termination of the feedback loop and thus increased the impact of the biological control agent on the target. The overall effectiveness of this classical biological control program can be attributed, in part, to the phenomenon of the target plant’s induced susceptible response to a herbivore.     

Chemotype variation of the weed Melaleuca quinquenervia influences the biomass and fecundity of the biological control agent Oxyops vitiosa

Host plant nutritional and non-nutritional variability can have a significant effect on herbivore populations by influencing survival, larval performance, and fecundity. The effect of chemical and physical variation of the leaves of two chemotypes of the weed Melaleuca quinquenervia was determined on the biomass and fecundity of the biological control agent Oxyops vitiosa (Coleoptera: Curculionidae). M. quinquenervia chemotypes were distinguished by the principal terpenoids E-nerolidol and viridiflorol using gas chromatography and mass spectroscopy. Not only were the terpenoid profiles of the two chemotypes different but the viridiflorol leaves had greater toughness (1.2-fold) and reduced nitrogen (0.7-fold). When the larvae and adults were fed leaves of the E-nerolidol chemotype increased adult biomass (1.1-fold) and fecundity were found (2.6- to 4.5-fold) compared with those fed leaves of the viridiflorol chemotype. Regardless of the larval diet, when adults were fed the E-nerolidol chemotype leaves they had greater egg production compared with those adults fed the viridiflorol leaves. Moreover, adult pre-oviposition period was extended (1.5-fold) when individuals were fed the viridiflorol leaves compared with those fed the E-nerolidol leaves. By rearing the O. vitiosa weevil on the more nutritious chemotype plants these results assisted in the mass production and establishment of the M. quinquenervia biological control agent.     

Maintenance of a narrow host range by Oxyops vitiosa; a biological control agent of Melaleuca quinquenervia

Host range expansion in insect herbivores is often thought to be mediated by several factors, principal among them are secondary plant metabolites. In weed biological control, the host range of a prospective agent is one of the most important considerations in its implementation. Extensive host testing tests seek to determine the behavioral acceptance and nutritional value of different test plant species to the potential agent. A list of test plants is compiled that comprises species that are close taxonomic relatives of the target weed plus other species of economic or ecologic importance. The host testing of the Melaleuca quinquenervia biological control agent Oxyops vitiosa indicated that larvae would accept and complete development on the Australian target weed M. quinquenervia, two Australian ornamental species, Callistemon citrina, Callistemon viminalis (all Myrtaceae). However, the larvae did not complete development when fed a North American species Myrica cerifera (Myricaceae). The study reported here confirms these results and examines the nutritional and performance differences in O. vitiosa larvae fed leaves of these species. The leaf quality factors, percent moisture, percent nitrogen, toughness, and terpenoid content were related to larval survival, performance and digestive indices. The results indicate that plant quality among the Myrtaceae species was generally similar and correspondingly larval survival, performance and digestive indices differed little when larvae were fed leaves of these species. However, significant differences occurred in the plant quality of the North American M. cerifera compared with the Australian species which had leaves with the lowest percent moisture, lowest leaf toughness, highest percent nitrogen. This species, however, is not a physiological host as none of the neonates survived to pupate. When third instars were switched to M. cerifera from their normal host M. quinquenervia reductions were found in survival, biomass gain, digestive efficiency, and conversion of digested food to insect biomass. The marginal acceptance of this North American native plant in laboratory bioassays appears related to the terpenoid chemistry that has similarities to the taxonomically unrelated host M. quinquenervia. However, the high larval mortality corresponds to several novel terpenoids that are not present in the host. For weed biological control host testing these results indicate that M. cerifera is a poor host for O. vitiosa. Additionally, future test plant lists should include plants with secondary metabolites similar to the target weed as these compounds may constitute behavioral cues that are relevant to these specialized herbivores.     

Host Plant Quality Factors That Influence the Growth and Development of Oxyops vitiosa, a Biological Control Agent of Melaleuca quinquenervia

The Australian weevil Oxyops vitiosa was released in 1997 in Florida as a biological control agent of Melaleuca quinquenervia. The larvae of this agent are flush-feeders, found only on the growing tips of their host. Knowledge of this restriction to feeding on the growing tips and other nutritional requirements may assist in the establishment and dispersal of this species. Therefore, O. vitiosa survival was assessed when neonates were fed M. quinquenervia leaves from branches that had dormant buds or emerging bud leaves. Additionally, the influence of leaf quality from different sites and within sites was determined by the feeding of neonates emerging bud leaves collected at three sites and from three leaf qualities (poor, intermediate, and high). Within-site leaf qualities were described in the field by leaf color and in the laboratory by percentage dry mass and nitrogen. Larval survival was lowest when fed leaves from branches that had dormant buds. Associated with this low survival were high leaf toughness and percentage dry mass. When larvae were fed emerging bud leaves, most of the variation in larval survival and performance was attributed to differences in within-site plant quality. Generally, the highest-quality leaves had relatively low percentage dry mass and high percentage nitrogen. Larval survival generally decreased when fed the poor-quality leaves, and in one site, the intermediate-quality leaves. Larvae required less time to develop to adults when fed the high-quality leaves. Development time increased in females but not in males when the larvae were fed the poor-quality leaves. Adult biomass of both females and males generally increased when the larvae were fed the high-quality leaves from two of the three sites. The results indicate that the larvae of O. vitiosa are restricted to feeding on flush foliage with low toughness. Additionally, variations in foliar percentage dry mass and nitrogen influence larval survival and performance. This knowledge benefited the development of mass-production nursery sites and the selection of suitable release sites, which facilitated the establishment of this biological control agent.     

The molecular basis of host plant selection in Melaleuca quinquenervia by a successful biological control agent

Melaleuca quinquenervia (Cav) S.T. Blake (broadleaf paperbark) is an Australian tree that has become a serious weed in many places around the world. Two insects Oxyops vitiosa (the melaleuca weevil), and Boreioglycaspis melaleucae (the melaleuca psyllid), which were introduced to Florida as part of a biological control programme, have been very effective in reducing survival and reproduction of this weed. There are two terpene chemotypes of M. quinquenervia; one rich in the sesquiterpene E-nerolidol whereas the other is rich in viridiflorol. Viridiflorol is a strong feeding deterrent for the melaleuca weevil and retards larval development. The larvae therefore avoid the viridiflorol-rich chemotype, in contrast, female melaleuca psyllids prefer to oviposit on these leaves. To identify the molecular basis of these preferences, we isolated and characterised two terpene synthases from the viridiflorol-rich chemotype, both of which utilise farnesyl pyrophosphate and have the same product profile. Chemotypic variation in terpenes in M. quinquenervia is under strong genetic control and the reproductive potential of each chemotype is limited by a different insect. These insects could, therefore, be selective agents for the maintenance of chemotypic variation in M. quinquenervia.     

Within-plant distribution of the Melaleuca quinquenervia biological control agent Lophodiplosis trifida

The gall-producing midge Lophodiplosis trifida Gagné (Diptera: Cecidomyiidae) is an established biological control agent of the exotic tree Melaleuca quinquenervia (Cav.) Blake, which invades wetland systems of Florida (USA). Host use patterns within the tree canopy were investigated. Plant height affected within-plant distribution of galls as L. trifida attack rates were greater in lower versus higher portions of the M. quinquenervia canopy. Nonetheless, galls occurred even at tree-top levels of 13 m.     

Invasion of a West Everglades wetland by Melaleuca quinquenervia countered by classical biological control

The population dynamics of Melaleuca quinquenervia were monitored over a 5-year period in a cypress-pine wetland while subjected to two levels of herbivory. The trees had been recruited during 1998–1999 after a destructive crown fire. Half of 26 experimental plots were sprayed every 4–6 weeks with an insecticide to reduce herbivory by the biological control agents Oxyops vitiosa and Boreioglycaspis melaleucae. After only 1-year melaleuca density increased by 26% in sprayed plots and by 7% in unsprayed plots. However, over the entire 5-year period melaleuca density increased in sprayed plots by 0.1% while decreasing by 47.9% in unsprayed plots when compared to initial densities. Annual mortality of melaleuca never exceeded 6% in any year in sprayed plots but ranged from 11% to 25% in unsprayed plots. There was a significant year by treatment interaction indicating the importance of the environment on tree mortality. Limited seed production occurred on sprayed trees but never on unsprayed trees. Mean tree height increased by 19.6% in sprayed plots while declining by 30.6% in unsprayed plots. Coverage by native vegetation did not increase with decreasing melaleuca density. This is the first study with controls that quantifies the population level regulation of melaleuca by introduced biological control agents and corroborates other correlative studies that documented significant changes in melaleuca communities after the introduction and establishment of biological control agents.     

An evaluation of the impact of Melaleuca quinquenervia invasion and management on plant community structure after fire

Two questions were asked in this study: after a fire, does the choice of invasive plant management strategy, namely herbicidal or biological, alter (1) plant community assemblages and (2) the re-invasion potential of the Australian tree Melaleuca quinquenervia? Plant species richness was highest in the non-invaded and herbicide sites compared to the biological site with 10.5, 10.8, and 8.25 species m⁻² found in each site, respectively. Although the total count of live and dead seedlings was highest in the biologically controlled site at 22.8 and 13.6 plants m⁻², respectively, M. quinquenervia seedlings were recruited in all sites. While the ultimate goal of management programs is to restore ecosystem integrity, this work provides evidence that passive restoration may not be enough to restore plant community structure in this system.     

Host range of Boreioglycaspis melaleucae Moore (Hemiptera: Psyllidae), a potential biocontrol agent of Melaleuca quinquenervia (Cav.) S.T. Blake (Myrtaceae), under quarantine

Research on the natural enemies of the Australian tree Melaleuca quinquenervia, a dominant, adventive weed of wetland habitats in south Florida, USA began in the late 1970s. Since then host-range tests have been conducted in Australia and Florida on three insect species: a weevil, a sawfly, and a psyllid. Herein we discuss the host-range studies of the psyllid, Boreioglycaspis melaleucae conducted in quarantine in Florida. Tests showed that the host range is very narrow, limited to one Melaleuca species, M. quinquenervia. One time, a few psyllids developed on a different species, Callistemon (=Melaleuca) citrinus, broad-leaf form, but repeated efforts to rear it again failed. In its Australian homeland, B. melaleucae has been collected on 5 closely related broad-leaved Melaleuca species including M. quinquenervia; it developed on 2 of 4 species tested, M. viridiflora (28%) and M. quinquenervia (37%). There was also 2% development on a 6th narrow-leaved Melaleuca species, M. nodosa. In the New World, there are no native Melaleuca species; in the United States, there is one introduced group, Callistemon (=Melaleuca), which is of commercial value and just one naturalized species, M. quinquenervia, which has no commercial value. Because of the near absence of Melaleuca species and B. melaleucae’s narrow host range and ability to severely damage or kill M. quinquenervia, we concluded that field release of B. melaleucae in south Florida would be safe. The first releases were made in February 2002.     

Post-biological control invasion trajectory for Melaleuca quinquenervia in a seasonally inundated wetland

The recruitment and mortality of Melaleuca quinquenervia seedlings were evaluated over a 3-year period in a seasonally inundated wetland in the western Everglades region. The mean (±SE) density of seedlings/saplings m^?1 declined from 64.8 (±4.5) to 0.5 (±0.2) over the 3 years, a population reduction of 99.2%. Four distinct water regimes characterized this site: dry, dry to wet transition, flooded, and wet to dry transition. Seedling recruitment was highest in the dry to wet transition and lowest in the flooded water regime, while mortality was highest under flooded and dry water regimes. The mean estimate of population growth (λ) across water regimes was 0.64 ± 0.05 indicating negative population growth. Elimination of introduced insect herbivores using insecticides did not reduce mortality of recruited M. quinquenervia seedlings/saplings indicating that direct herbivory was not responsible for the decline in seedling density. On the other hand, a mean of only 0.2 (±0.03) viable seeds m^?2 d^?1 fell into the plots, an amount considerably lower than in previous studies. We submit that change in the invasion trajectory M. quinquenervia was most likely caused by reduced seed inputs from aerial seed banks depleted by insect herbivory rather than direct herbivory on seedlings. This may indicate a fundamental alteration of M. quinquenervia population dynamics ultimately resulting in a less invasive and, therefore, less ecologically damaging species.     

Carbon isotope discrimination in leaves of the broad‐leaved paperbark tree,Melaleuca quinquenervia,as a tool for quantifying past tropical and subtropical rainfall

Quantitative reconstructions of terrestrial climate are highly sought after but rare, particularly in Australia. Carbon isotope discrimination in plant leaves (Δ_leaf) is an established indicator of past hydroclimate because the fractionation of carbon isotopes during photosynthesis is strongly influenced by water stress. Leaves of the evergreen tree Melaleuca quinquenervia have been recovered from the sediments of some perched lakes on North Stradbroke and Fraser Islands, south‐east Queensland, eastern Australia. Here, we examine the potential for using M. quinquenervia ?_leaf as a tracer of past rainfall by analysing carbon isotope ratios (δ¹³C) of modern leaves. We firstly assess Δ_leaf variation at the leaf and stand scale and find no systematic pattern within leaves or between leaves due to their position on the tree. We then examine the relationships between climate and Δ_leaf for a 11‐year time series of leaves collected in a litter tray. M. quinquenervia retains its leaves for 1–4 years; thus, cumulative average climate data are used. There is a significant relationship between annual mean ?_leaf and mean annual rainfall of the hydrological year for 1–4 years (i.e. 365–1460 days) prior to leaf fall (r² = 0.64, P = 0.003, n = 11). This relationship is marginally improved by accounting for the effect of pCO₂ on discrimination (r² = 0.67, P = 0.002, n = 11). The correlation between rainfall and Δ_leaf, and the natural distribution of Melaleuca quinquenervia around wetlands of eastern Australia, Papua New Guinea and New Caledonia offers significant potential to infer past rainfall on a wide range of spatial and temporal scales.     

Decline in exotic tree density facilitates increased plant diversity: the experience from Melaleuca quinquenervia invaded wetlands

The Australian tree Melaleuca quinquenervia (melaleuca) formed dense monocultural forests several decades after invading parts of Florida and the Caribbean islands. These dominant forests have displaced native vegetation in sensitive wetland systems. We hypothesized that native plant diversity would increase following recent reductions in density of mature melaleuca stands in south Florida. We therefore examined data on changes in melaleuca densities and plant species diversity derived from permanent plots that were monitored from 1997 to 2005. These plots were located within mature melaleuca stands in nonflooded and seasonally-flooded habitats. Two host-specific biological control agents of melaleuca, Oxyops vitiosa and Boreioglycaspis melaleucae, were introduced during 1997 and 2002, respectively. Also, an adventive rust fungus Puccinia psidii and lobate-lac scale Paratachardina pesudolobata became abundant during the latter part of the study period. Overall melaleuca density declines in current study coincided with two to four fold increases in plant species diversity. The greatest declines in melaleuca density as well as the greatest increases in family importance values and species diversity indices occurred in nonflooded as compared to seasonally-flooded habitats. Most pioneer plant species in study sites belonged to Asteraceae, Cyperaceae, Poaceae, and Ulmaceae. The rapid reduction in melaleuca density and canopy cover during the study period may be attributed to self-thinning accelerated by the negative impact of natural enemies. Densities of other woody plants, particularly Myrica and Myrsine, which were sparsely represented in the understory by a few suppressed individuals also declined during the same period, possibly due to infestation by the generalist lac-scale. These findings indicate that natural-enemy accelerated self-thinning of melaleuca densities is positively influencing the native plant diversity and facilitating the partial rehabilitation of degraded habitats.     

Activity rhythms of the shallow-water octopuses Octopus laqueus and Abdopus aculeatus with special reference to its relation to light and tidal cycles

Patterns of activity and inactivity were experimentally measured for two shallow-water octopuses, Octopus laqueus (n = 8) and Abdopus aculeatus (n = 4), inhabiting the Ryukyu Archipelago. Octopuses that were collected from the coastal waters of Okinawa Island were held in experimental tanks under controlled light conditions (L, light; D, dark; 12L:12D, LD conditions; 12D:12D, DD conditions). Behaviors of these two species were continuously recorded for 9 to 10 days. Under LD conditions, O. laqueus were active for 7 to 14 min during daytime and 298 to 339 min at night, and under DD conditions, for 97 to 140 min during daytime and 71 to 169 min at night. A. aculeatus were active for 49 to 99 min during daytime and 138 to 185 min at night under LD conditions, whereas under DD conditions, they were active for 36 to 56 min during daytime and 55 to 154 min at night. Continuous duration for activity cycles was 39 ± 7.6 h under LD conditions and 42 ± 2.6 h under DD conditions in O. laqueus, and 38 ± 7.4 h under LD conditions and 42 ± 2.8 h under DD conditions in A. aculeatus. Ratios for active duration to inactive duration did not differ between LD condition and DD condition both in O. laqueus and A. aculeatus.     

Physiological host range of two highly specialised mutualistic symbiotes: the fly Fergusonina turneri and its obligate nematode Fergusobia quinquenerviae,potential biocontrol agents of Melaleuca quinquenervia

In Australia, galls develop on Melaleuca quinquenervia (Cav.) S.T. Blake (Myrtaceae) as a result of the mutualistic association between the fly Fergusonina turneri Taylor (Diptera: Fergusoninidae) and its obligate nematode Fergusobia quinquenerviae Davies & Giblin-Davis (Tylenchida: Sphaerulariidae). The nematode induces gall formation, whereas the fly promotes gall maturation. Together they exploit M. quinquenervia buds and may inhibit stem elongation and flower formation. We delimited the physiological host range of this pair to determine their suitability as biological control agents of invasive M. quinquenervia populations in Florida, USA. Host use was assessed for eight species of Myrtaceae native to Florida, eight phylogenetically related ornamental species and oviposition alone on five non-myrtaceous species. Although oviposition was less specific, galls developed and matured only on M. quinquenervia. After establishment, galls are predicted to prevent flower and seed production, thereby reducing the regenerative potential of M. quinquenervia. This is the first example of an insect/nematode mutualism released as biological control agents of an invasive plant.