Identification of the Klambothrips myopori (Thysanoptera: Phlaeothripidae) Predator Complex in California.

To identify the predator complex of the invasive thrips, Klambothrips myopori, on its ornamental host plant Myoporum laetum, field surveys were conducted at three sites in southern California over the period of 1 y. Five insect orders and five spider families were represented in the survey. Although the most abundant groups differed among collection sites, syrphid larvae, anthocorids, Chrysoperla spp., Franklinothrips orizabensis Johansen (Thysanoptera: Aeolothripidae), and one spider family (Salticidae) were all collected at each site. Based on the field surveys, Orius spp. and Chrysoperla spp. were identified as possible key natural enemies of K. myopori. Laboratory studies were then conducted to determine the consumption rates of Orius insidiosus Say (Hemiptera: Anthocoridae) and Chrysoperla rufilabris Burmeister (Neuroptera: Chrysopidae) at constant densities of K. myopori and to define the functional responses of the predators. Both predators consumed more second-instar larvae than other prey stages. Orius insidious displayed a type II functional response, while C. rufilabris displayed both type II and type III depending on prey stage. Generally speaking, O. insidiosus and C. rufilabris consumed a higher proportion of prey at lower pest densities, implying that in an augmentative control program using these commercially available natural enemies, predators could be released early in the year when host plants begin to flush and thrips populations are low to suppress population growth.

Economic Benefit for Cuban Laurel Thrips Biological Control.

The Cuban laurel thrips, Gynaikothrips ficorum Marchal (Thysanoptera: Phlaeothripidae), is a critical insect pest of Ficus microcarpa in California urban landscapes and production nurseries. Female thrips feed and oviposit on young Ficus leaves, causing the expanding leaves to fold or curl into a discolored leaf gall. There have been attempts to establish specialist predator natural enemies of the thrips, but no success has been reported. We resampled the same areas in 2013-2014 where we had released Montandoniola confusa (= morguesi) Streito and Matocq (Hemiptera: Anthocoridae) in southern California in 1995 but had been unable to recover individuals in 1997-1998. Thrips galls were significantly reduced in all three of the locations in the recent samples compared with the earlier samples. M. confusa was present in all locations and appears to be providing successful biological control. The value of the biological control, the difference between street trees in good foliage condition and trees with poor foliage, was $58,766,166. If thrips damage reduced the foliage to very poor condition, the value of biological control was $73,402,683. Total cost for the project was $61,830. The benefit accrued for every dollar spent on the biological control of the thrips ranged from $950, if the foliage was in poor condition, to $1,187, if the foliage was in very poor condition. The value of urban forest is often underappreciated. Economic analyses that clearly demonstrate the very substantial rates of return on investment in successful biological control in urban forests provide compelling arguments for supporting future efforts.

Cost-Benefit Analysis for Biological Control Programs That Targeted Insect Pests of Eucalypts in Urban Landscapes of California.

As well as being planted for wind breaks, landscape trees, and fuel wood, eucalypts are also widely used as urban street trees in California. They now are besieged by exotic insect herbivores of four different feeding guilds. The objective of the current analysis was to determine the return on investment from biological control programs that have targeted these pests. Independent estimates of the total number of eucalypt street trees in California ranged from a high of 476,527 trees (based on tree inventories from 135 California cities) to a low of 190,666 trees (based on 49 tree inventories). Based on a survey of 3,512 trees, the estimated mean value of an individual eucalypt was US$5,978. Thus, the total value of eucalypt street trees in California ranged from more than US$1.0 billion to more than US$2.8 billion. Biological control programs that targeted pests of eucalypts in California have cost US$2,663,097 in extramural grants and University of California salaries. Consequently, the return derived from protecting the value of this resource through the biological control efforts, per dollar expended, ranged from US$1,070 for the high estimated number of trees to US$428 for the lower estimate. The analyses demonstrate both the tremendous value of urban street trees, and the benefits that stem from successful biological control programs aimed at preserving these trees. Economic analyses such as this, which demonstrate the substantial rates of return from successful biological control of invasive pests, may play a key role in developing both grass-roots and governmental support for future urban biological control efforts.

Potential for Interspecific Competition Between Congeneric Longhorned Beetle Species (Coleoptera: Cerambycidae) in an Adventive Environment.

The cerambycid beetle, Phoracantha semipunctata F., was introduced into California in the mid-1980s and killed large numbers of Eucalyptus host trees. The populations of the borer declined to very low levels in the mid-1990s following the establishment of the congener, Phoracantha recurva Newman, and the intentional introduction of the egg parasitoid, Avetianella longoi Siscaro. The distributions of the beetles overlap in the Australian native range, but one species has replaced the other in the adventive range in California. One possible explanation is differential susceptibility to natural enemies introduced for biological control. An alternative explanation for the reduced abundance of P. semipunctata is asymmetric interspecific competition between the two species. To test this hypothesis, equal larval densities of each species were introduced into host logs. In all cases, more P. recurva adults emerged than P. semipunctata adults, but the presence of congeners did not have a different effect than the presence of an equal density of conspecific individuals. Neither the temporal order of introduction or bark thickness altered the outcome of potential competitive interactions. Consequently, it appears that the ecological replacement of one borer with another in the adventive environment in southern California may not be a result of bottom-up intraguild competitive interactions. The top-down effects of natural enemies on P. semipunctata have most likely led to its decline.

A new Huanglongbing Species, "Candidatus Liberibacter psyllaurous," found to infect tomato and potato, is vectored by the psyllid Bactericera cockerelli (Sulc).

A new huanglongbing (HLB) "Candidatus Liberibacter" species is genetically characterized, and the bacterium is designated "Candidatus Liberibacter psyllaurous." This bacterium infects the psyllid Bactericera cockerelli and its solanaceous host plants potato and tomato, potentially resulting in "psyllid yellowing." Host plant-dependent HLB transmission and variation in psyllid infection frequencies are found.

Frequency of secondary symbiont infection in an invasive psyllid relates to parasitism pressure on a geographic scale in California.

Two endosymbionts, an obligate primary symbiont and a facultative secondary symbiont, are harbored within the invasive red gum (eucalyptus) lerp psyllid, Glycaspis brimblecombei, in California. An extensive survey of diversity and frequency of G. brimblecombei's secondary symbiont in multiple populations throughout the state of California was conducted using PCR detection, restriction enzymes, cloning, and sequencing. A total of 380 G. brimblecombei individuals in 19 populations were screened for secondary symbionts. Based on molecular screening results, only one type of secondary symbiont was present in G. brimblecombei populations in California. Overall, 40% of the 380 psyllids screened were infected with the secondary symbiont. Interestingly, secondary symbiont infection frequencies in G. brimblecombei populations varied dramatically from 0 to 75% and were significantly related to parasitism pressure by Psyllaphaegus bliteus, a solitary endoparasitoid of the psyllid.

Aggressive interactions between Solenopsis invicta and Linepithema humile (Hymenoptera: Formicidae) under laboratory conditions.

The Argentine ant, Linepithema humile (Mayr), and the red imported fire ant, Solenopsis invicta Buren, are natural agonists in their country of origin. Since the first report of L. humile in California in 1907 its range expanded statewide, displacing native ant species wherever it spread. Since the discovery of established populations of S. invicta in southern California in 1998, it has been restricted to discrete areas of southern California. However, as these discrete populations expand, they are encountering large populations of L. humile, which are effective competitors for available resources and are particularly aggressive in their encounters with other ant species such as S. invicta. Most Dolichoderine ants such as L. humile do not prefer to forage on baits made with defatted corn grit and soybean oil typically used in red imported fire ant control programs. Applications of these baits in areas where distributions of these species overlap might selectively affect populations of S. invicta and give L. humile a competitive advantage. Three laboratory experiments were conducted to determine the competitive outcomes between S. invicta pitted against L. humile: 1) agonistic behavior of workers in small arenas, 2) colony interactions with different population ratios, and 3) the effects of pyriproxyfen on the competitiveness of S. invicta against L. humile. Populations of S. invicta consisting of major workers killed more L. humile than did minors or a mixture of majors and minors. When paired against L. humile colonies consisting of 1,100 workers, colonies consisting of 38 S. invicta workers were easily defeated by L. humile. Colonies consisting of 450 S. invicta workers plugged their nest entrances, but they were ultimately defeated by L. humile after 13 d. The S. invicta colonies consisting of 1,100 workers took control of the bridge connecting the colonies, invaded the L. humile colony, killed the Argentine ant queens, and removed their brood. Pyriproxyfen-treated fire ant workers took significantly longer to chase the Argentine ants from a connecting bridge than did untreated fire ants. Thus, fire ant baits may have long-term effects on intercolonial aggression between S. invicta and L. humile, especially when Argentine ant populations are high in the summer.

Genetic relationships among Leptographium terebrantis and the mycangial fungi of three western Dendroctonus bark beetles.

Morphology, mitochondrial DNA (mtDNA) restriction fragment polymorphisms (RFLPs) and nuclear DNA (nDNA) fingerprinting were used to clarify relationships among the morphologically similar Ophiostoma and Leptographium species associated with mycangia of three Dendroctonus bark beetles (Ophiostoma clavigerum associated with both D. ponderosae and D. jeffreyi, and L. pyrinum associated with D. adjunctus), as well as a closely related nonmycangial bark beetle associate (L. terebrantis). Most isolates of O. clavigerum form long (40-70 µm), septate conidia, while all isolates of L. terebrantis and L. pyrinum form conidia less than 17.0 µm in length. The conidia of L. pyrinum are pyriform, with truncate bases, while the conidia of the other species form only slightly truncate bases. Conidial masses of L. terebrantis are creamy yellow, while the conidial masses of the other species are white. Nuclear DNA fingerprints resulting from probing PstI restrictions with the oligonucleotide probe (CAC)(5) and HaeIII and MspI restrictions of mtDNA, exhibited three major clusters. In the dendrogram developed from mtDNA RFLPs, the L. pyrinum isolates formed one cluster, while the majority of O. clavigerum isolates, including all D. jeffreyi isolates, formed another. A third cluster was composed of all L. terebrantis isolates, as well as several O. clavigerum isolates from D. ponderosae. The inclusion of some O. clavigerum isolates in the L. terebrantis cluster suggests that horizontal transfer of mtDNA has occurred among these fungi. The nDNA dendrogram also exhibited three clusters, and most isolates of L. pyrinum, L. terebrantis and O. clavigerum grouped separately; however, one isolate of O. clavigerum grouped with the L. terebrantis isolates, while one isolate of L. terebrantis grouped with O. clavigerum. No genetic markers were found that distinguished between O. clavigerum associated with D. ponderosae and O. clavigerum associated with D. jeffreyi. Ophiostoma clavigerum might be a recently diverged morphological variant of L. terebrantis, with special adaptations for grazing by young adults of D. jeffreyi and D. ponderosae. The anamorph of O. clavigerum, Graphiocladiella clavigerum, is transferred to Leptographium.

Methods for rearing Syngaster lepidus and Jarra phoracantha (Hymenoptera: Braconidae), larval parasitoids of the phloem-colonizing longhorned beetles Phoracantha semipunctata and P. recurva (Coleoptera: Cerambycidae).

Two species of cerambycid beetles that attack eucalypts, Phoracantha semipunctata (Fabricius) and P. recurva Newman, have been accidentally introduced from Australia into most regions of the world in which their hosts have been planted. The beetles cause extensive mortality in plantations and landscape plantings of the trees. Management programmes have focused on integration of silvicultural practices, host plant resistance and biological control. To rear and release natural enemies of the larval stages of the beetles in California, mass rearing protocols for continuous production of two species of parasitoids have been developed. The methods described represent the first long-term and large-scale techniques for mass rearing parasitoids of any wood-boring cerambycid species. In addition to providing large numbers of parasitoids for releases, the mass rearing effort has also provided large numbers of parasitoids for fundamental studies of their biology and behaviour.

Insect pests of eucalypts in California: implications of managing invasive species.

For the first 150 years following their introduction, eucalypts planted in the California landscape were free of both insect pests and diseases. In the last 15 years, numerous herbivorous insect species have been introduced accidentally into the State and have caused significant damage to the trees. Several of these species, e.g. Phoracantha semipunctata (Fabricius), Phoracantha recurva Newman (Coleoptera: Cerambycidae) and Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), have also been introduced into other parts of the world where eucalypts are grown, whereas others, e.g. Glycaspsis brimblecombei Moore (Hemiptera: Spondyliaspidae) and Eucalyptolyma maideni Froggatt (Hemiptera: Spondyliaspidae), are currently restricted to California and Australia. Research programmes have provided management solutions to individual pest problems, but as more pest species are introduced, these solutions must be integrated across broad geographic, horticultural, and economic scales, in a systems approach.

Interactions among Scolytid bark beetles, their associated fungi, and live host conifers.

Scolytid bark beetles that colonize living conifers are frequently associated with specific fungi that are carried in specialized structures or on the body surface. These fungi are introduced into the tree during the attack process. The continuing association suggests that there is mutual benefit to the fitness of both beetles and fungi. The fungal species may benefit from the association with the beetles by transport to new host trees. Beetle species may benefit from the association with fungi by feeding on the fungi, or by the fungi contributing to the death of the host trees through mycelial penetration of host tissue, toxin release, interactions with preformed and induced conifer defenses, or the combined action of both beetles and fungi during colonization. Extensive research has been directed towards characterizing the interactions of beetle-fungal complexes with live host conifers and determining the ecological advantages for maintaining the associations. However, differences among systems and how species interact under different population and environmental conditions make it difficult to generalize about the importance of the separate biological components in successful host colonization.

Interactions between the Encelia leaf beetle and its host plant, Encelia farinosa: The influence of acidic fog on insect growth and plant chemistry.

The impact of acidic deposition on interactions between the plant Encelia farinosa and the herbivorous beetle Trirhabda geminata (Chrysomelidae) was determined under greenhouse conditions. Acidic fogs (pH 2.75) did not significantly affect the overall foliar concentrations of water or soluble protein as compared with control fogs (pH 5.6). Nonetheless, E. farinosa foliage was altered by exposure to three 3-h acidic fogs such that growth and biomass gain by T. geminata increased by more than 30% as compared to beetles feeding on control-fogged plants. Thus, previous indications that changes in soluble proteins or water content were responsible for increased biomass gain and growth of T. geminata cannot be substantiated by this study. Additionally, changes in the plant defensive chemistry were not responsible for increased herbivore growth, as farinosin, encecalin, and euparin foliar concentrations did not vary significantly between fog treatments. Significant increases in CO2 assimilation rates of E. farinosa exposed to acidic fogs were documented at 3, 7, and 21 days following treatment, suggesting that carbohydrate-based products of increased plant metabolism may have played a role (e.g. soluble carbohydrates). However, the key factors responsible for increasing herbivore performance on acidic-fogged E. farinosa remain largely unknown.

Cardiac transplant coronary artery disease. A multivariable analysis of pretransplantation risk factors for disease development and morbid events.

Coronary artery disease after cardiac transplantation is a major obstacle to long-term survival. The development and progression of coronary artery disease after cardiac transplantation was analyzed in 217 consecutive patients undergoing transplantation. The actuarial freedom from any coronary artery disease (by angiography or autopsy) was 81% at 2 years and 20% at 8 years after transplantation. Coronary artery disease was more prevalent in male than female patients (30% versus 50% free of coronary artery disease at 5 years, p = 0.01). By multivariable analysis, pretransplantation risk factors identified for coronary artery disease included pretransplantation positive cytomegalovirus serologic status of the recipient (p = 0.002) and older donor age (p = 0.07). Progression of coronary artery disease was variable in both time of onset and rate. Earlier detection did not result in more rapid progression. Coronary events severe enough for retransplantation (n = 8) and/or death from coronary artery disease (n = 9) occurred in 15 patients, of whom four underwent retransplantation. The actuarial freedom from coronary events was 88% at 5 years and 79% at 8 years. By multivariable analysis, only male recipient (p = 0.05) was a risk factor for coronary events. Seven of the 15 patients (47%) with coronary events died suddenly of coronary artery disease without prior angiographic evidence of severe coronary disease. Coronary artery disease is progressive. Improved surveillance methods are required to detect the disease and institute timely intervention to prevent the occurrence of unanticipated death.

Trirhabda geminata (Coleoptera: Chrysomelidae) resistance to the direct impact of simulated acidic fog on larval growth and mortality.

The direct effects of acidic fog (pH 2.75) upon the mortality and growth of Trirhabda geminata Horn (Coleoptera: Chrysomelidae), the dominant herbivore of the California coastal sage scrub, Encelia farinosa (Compositae: Asteraceae), were evaluated. Although there was a consistent pattern among and between experiments, suggesting that acidic fogs could reduce insect growth and survivorship within the first few days following application of treatments, an exposure to three consecutive, 3-h fogs over a five day period did not significantly affect mortality, biomass gain, or larval growth rate. There are two important implications from this study. First, even the highly acidic fogs found in southern California will have minimal direct effects on T. geminata performance. Second, the impacts on T. geminata biology observed in previous studies were likely mediated by host-plant responses to acidic-fog episodes.

Influence of oleoresin constituents fromPinus ponderosa andPinus jeffreyi on growth of mycangial fungi fromDendroctonus ponderosae andDendroctonus jeffreyi.

Dendroctonus jeffreyi andD. ponderosae are morphologically similar sympatric species of pine bark beetles over portions of their geographic ranges; however,D. jeffreyi is monophagous onP. jeffreyi whileD. ponderosae is highly polyphagous. Both species carry a species of mycangial fungi that are also very similar in appearance. Growth of the two mycangial fungi and of the fungusLeptographium terebrantis (associated with the polyphagous and non-tree-killingDendroctonus valens) in the presence of oleoresin constituents of host and nonhost conifers was tested by placing individual chemicals on agar growth medium and by growing the cultures in saturated atmospheres of the chemicals. The fungus associated withD. jeffreyi showed greater tolerance for chemical constituents placed on the growth medium than the other two fungi, and growth after three days was enhanced by heptane, the dominant constituent ofP. jeffreyi oleoresin. Growth of all three species of fungi was reduced by the resin constituents when the chemicals were presented as saturated atmospheres. The results suggest that the influence of the tree on growth of the symbiotic fungi of the bark beetles during the initial attack process may be different than after colonization by the beetles is complete. The difference in the responses of the apparently related species of mycangial fungi may provide some new insight into the evolutionary history of these beetle/mycangial fungus/host tree systems.

Response ofDendroctonus brevicomis Le Conte (Coleoptera: Scolytidae) to different release rates and ratios of aggregation semiochemicals and the inhibitors verbenone and ipsdienol.

The response of western pine beetle,Dendroctonus brevicomis Le Conte, to different release rates of the aggregation semiochemicals (attractants)exo-brevicomin, frontalin, and myrcene and the inhibitors verbenone and ipsdienol was investigated. Release rates of verbenone ranging from 0.18 mg/ 24 hr to 1.2 mg/24 hr did not result in significant reductions in mean trap catch ofD. brevicomis. In contrast, very low release rates of ipsdienol (0.02-0.4 mg/24 hr) significantly reduced trap catch compared to controls. The combination of verbenone and ipsdienol, released at rates above 0.09 and 0.02 mg/24 hr, respectively, resulted in significantly lower trap catches ofD. brevicomis in attractant-baited traps. Results of an experiment testing a factorial combination of different release rates of verbenone and attractants suggest that response is not ratio-specific. The response curve ofD. brevicomis to the levels of verbenone was similar across all levels of attractants, while the response to equivalent ratios of attractants to verbenone was not similar, suggesting that the behavior of the beetles is primarily influenced by the absolute release rate of verbenone.

Influence of aggregation inhibitors (verbenone and ipsidenol) on landing and attack behavior ofDendroctonus brevicomis (Coleoptera: Scolytidae).

The influence of the aggregation inhibitors verbenone and ipsdienol on the response of western pine beetle,Dendroctonus brevicomis, to attractive host trees was investigated. Paired ponderosa pine trees (Pinus ponderosa) were baited with aggregation semiochemicals to stimulate mass attack. One tree in each pair received an inhibitor treatment consisting of five sets of two verbenone and two ipsdienol dispensers spaced 1 m apart vertically along the tree bole. Beetle landing was monitored with sticky traps on the tree bole, and attack density was assessed from bark samples removed four or seven days after baiting. The inhibitor treatment resulted in a significant reduction of both the numbers of beetles landing on trees and the density of attacking beetles compared to control trees (without inhibitors). The ratios of beetle landing density to attacking density were not different between inhibitor-treated and control trees, nor were the vertical distributions of beetles landing or attacking, suggesting that beetle behavior was primarily influenced at a longer range, prior to landing on the tree. Although the application of verbenone and ipsdienol did not preventD. brevicomis from attacking baited trees, our results suggest that when applied to unattacked (and unbaited) trees, their effectiveness at reducing the attack pressure might allow trees having a certain amount of resistance to survive attack by pioneer beetles.

Impact of acidic deposition onEncelia farinosa gray (Compositae: Asteraceae) and feeding preferences ofTrirhabda geminata horn (Coleoptera: Chrysomelidae).

Container grownEncelia farinosa were exposed to three 3-hr episodes of acidic fog (pH 2.5) typical of events in southern California. Adults and larvae of the specialist leaf-feeding herbivore,Trirhabda geminata, preferred to feed on the acidic-treated foliage compared to control fogged (pH 6.3-6.5) foliage. Previous feeding damage on the plants did not affect feeding preference. The acidic-fogged foliage was significantly higher in total nitrogen and soluble protein but not different from control-treated tissue in water content. Stress on native populations of this drought-deciduous shrub caused by atmospheric pollutants may also result in altered feeding ecology of the beetle.

Control of the straw itch mite (Acari: Pyemotidae) with sulfur in an insect rearing facility.

The ectoparasitic mite Pyemotes tritici (Lagrèze-Fossat & Montané) (Acari: Pyemotidae) caused paralysis and reduced longevity in eucalyptus longhorned borer, Phoracantha semipunctata F., under laboratory rearing conditions. Application of dusting sulfur to logs that contained pupating borers greatly reduced densities of mites on emerging adult beetles and increased beetle survivorship. Uniform application to all logs in a glasshouse effectively eradicated the mite infestation. A bioassay showed that sulfur may physically impede the dispersal of immature mites by adhering to the cuticle, but sulfur vapor did not act as a toxin.

Response ofDendroctonus brevicomis andIps paraconfusus (Coleoptera: Scolytidae) to combinations of synthetic pheromone attractants and inhibitors verbenone and ipsdienol.

DendroctonusBrevicomis andIps paraconfusus are sympatric bark beetle species colonizingPinus ponderosa in western North America. Interspecific and intraspecific competition for resources is, in part, mediated through semiochemicals. The response ofD. brevicomis to its attractant pheromone was significantly reduced by simultaneous release of either verbenone or racemic ipsdienol. Trap catch was significantly further reduced by including both inhibitors with the attractant pheromones. However, although the response ofIps paraconfusus was significantly inhibited with the addition of either verbenone or racemic ipsdienol, both compounds together did not result in a significant further reduction in trap catch. There was a trend for greater reduction in response ofD. brevicomis to attractant pheromones with increased release rates of either 69% (+)-/31% (-)-verbenone or 84% (-)-/16% (+)-verbenone. Response of associates to attractants and inhibitor combinations was also determined.