The number of public-sector biological control scientists and practitioners in the United States was determined by a survey conducted in 2019. A total of 344 personnel were identified in 49 states and Washington DC; 218 employed by universities, 86 by the federal government and 36 by state departments of agriculture. There were 10-34 personnel in eight states, 6-9 in 11, and 0-5 in 31, overall averaging less than 50% of their combined effort on biological control. Applications for biological control agents included about 30% for row crops and vegetables; 12% each for orchards, urban landscapes, outdoor ornamentals, and forests; and 22% for the remainder. The federal government provided an average of 42% of the funding for university, federal, and state biological control research and extension. The states contributed 29-35%, commodity groups supplied 19-24%, and the remaining funds were from private sources. Of 54 universities in the United States, 18 conducted in-person classroom education and training in biological control, with seven offering courses via distance. However, another 18 universities had discontinued their comprehensive courses. Biological control instruction was incorporated into courses and workshops on integrated pest management and sustainable agriculture at 50 universities. Correspondingly, during the past 24 years, biological control has gradually developed from primarily an independent discipline to increasingly being part of integrated pest management. Classical, augmentative and conservation biological control have nevertheless remained distinguishable high priority subdisciplines. Funding continues to be essential for advancing biological control, requiring societal understanding and acceptance. Advancement of biological control in the United States will require increased investment in personnel and their education and training, along with delivery of effective pest management technologies.
Understanding and predicting potential competitive outcomes is critical in the design of biological control programs when considering multiple agent introductions. Tephritid fruit flies (Diptera: Tephritidae) include some of the most destructive pests of fruit and vegetable crops worldwide. Parasitoid guilds of fruit-infesting tephritids are dominated by opiines (Hymenoptera: Braconidae: Opiinae) that have been the main agents used for biological control of pest tephritids. All opiine parasitoids are solitary koinobiont endoparasitoids that attack host eggs or larvae and emerge as adults from host puparia. Thus, a host initially attacked by an egg-larval parasitoid can be subsequently attacked by a larval parasitoid, providing an ideal system to test the Early-acting Competitive Superiority Hypothesis (i.e., the first species occupying a host or exploiting early host stages has an advantage over competitive species). We reviewed the literature on interspecific competition among opiine fruit fly parasitoids, which reveals strong evidence supporting the early-acting competitive superiority hypothesis by egg-larval over larval parasitoids, through physiological suppression mechanisms. Competitive outcomes among larval parasitoids, however, depend on early action as well as the morphological characteristics used in direct physical competition of their first instars (e.g., size of mandibles). We discuss the range of ecological mechanisms facilitating coexistence of interacting species and highlight the ecological consequences of interspecific competition as evidenced by historical competitive displacements of introduced opiine parasitoids for biocontrol of tephritids. Finally, we provide a framework for exploring the role of interspecific competition, among other factors, in selecting opiine parasitoids for biocontrol of invasive tephritids.
The olive psyllid, Euphyllura olivina, is a newly invasive species to California with the potential to become an economical pest if it reaches the olive production regions of California's Central Valley. Here, we report on surveys undertaken in California to assess the psyllid's current distribution and the occurrence of parasitism. Additionally, we present results of foreign collections of its parasitoids and initial non-target studies of a possible biological control agent, the Mediterranean parasitoid Psyllaephagus euphyllurae. The current distribution of the psyllid appears to be limited to the California coast between Monterey and San Diego; there have been no reports of infestations on olives in the major production areas of central and northern California. Psyllaephagus euphyllurae was the major primary parasitoid found in our foreign collections. The potential non-target impact of P. euphyllurae was tested on three native North American psyllid species: Neophyllura arctostaphyli, Euglyptoneura nr. robusta, and Calophya nigrella. No P. euphyllurae developed on the non-target species during no-choice tests. Behavioral observations in choice tests confirmed no attack on the non-target hosts, although the parasitoid did remain longer on N. arctostaphyli-infested manzanita plants, and revealed no host feeding behavior.
Many invertebrate biological control agents (BCAs) and a few microbial pesticides are available to manage tomato pests, but considerably more that have been discovered and evaluated at some level could be produced commercially. To encourage further development of commercial products, most of the major arthropod pests of tomato are listed, along with some of their existing and potential BCAs and microbial pesticides. Biological control of tomato pests in protected culture and the field is reviewed, including sources of BCAs, quality assurance in their production, tomato grower evaluation of the products, and examples of their uses and costs. The sources and applications for microbial pesticides are also described. Expansion in the development and use of BCAs will require additional testing in tomato production systems, as will the generation of new microbial pesticide products. Advancements are needed in research and technology, regulatory oversight, and grower education to foster continued growth of commercial biological control for arthropod pests of tomato.
The establishment of olive fruit fly Bactrocera oleae (Rossi) in California, USA instigated a classical biological program. This study reports the release and recovery of two solitary larval endoparasitoids, Psyttalia humilis Silvestri and Psyttalia lounsburyi (Silvestri) imported from sub-Saharan Africa, and released in five coastal and three inland counties in California, USA. Both parasitoid species were recovered post-release within the same fruit season and dispersed up to 1,500 m from a release location. P. lounsburyi was recovered post-release the following fruit seasons at numerous sites, and up to 945 days after the last release at one site. It is now established in California coastal regions, but not at the inland release sites. We discuss ecological factors and aspects of parasitoid biology that could improve or impede the permanent establishment of olive fruit fly parasitoids in California.
Olive (Olea europaea) trees exhibiting leaf scorch or branch dieback symptoms in California were surveyed for the xylem-limited, fastidious bacterium Xylella fastidiosa. Only approximately 17% of diseased trees tested positive for X. fastidiosa by polymerase chain reaction, and disease symptoms could not be attributed to X. fastidiosa infection of olive in greenhouse pathogenicity assays. Six strains of X. fastidiosa were isolated from olive in Southern California. Molecular assays identified strains recovered from olive as belonging to X. fastidiosa subsp. multiplex. Pathogenicity testing of olive strains on grapevine and almond confirmed that X. fastidiosa strains isolated from olive yield disease phenotypes on almond and grapevine typical of those expected for subsp. multiplex. Mechanical inoculation of X. fastidiosa olive strains to olive resulted in infection at low efficiency but infections remained asymptomatic and tended to be self-limiting. Vector transmission assays demonstrated that glassy-winged sharpshooter (Homalodisca vitripennis) could transmit strains of both subspp. multiplex and fastidiosa to olive at low efficiency. Insect trapping data indicated that two vectors of X. fastidiosa, glassy-winged sharpshooter and green sharpshooter (Draeculacephala minerva), were active in olive orchards. Collectively, the data indicate that X. fastidiosa did not cause olive leaf scorch or branch dieback but olive may contribute to the epidemiology of X. fastidiosa-elicited diseases in California. Olive may serve as an alternative, albeit suboptimal, host of X. fastidiosa. Olive also may be a refuge where sharpshooter vectors evade intensive areawide insecticide treatment of citrus, the primary control method used in California to limit glassy-winged sharpshooter populations and, indirectly, epidemics of Pierce's disease of grapevine.
The overwintering survival and development of olive fruit fly, Bactrocera oleae (Rossi), and the endoparasitoids, Psyttalia humilis Silvestri and P. lounsburyi (Silvestri), were investigated at sites in California's interior valley and coastal region. In the interior valley, adult flies survived up to 4-6 mo during the winter when food was provided. Adult female flies could oviposit in late fall and early winter on nonharvested fruit and, although egg survival was low (0.23-8.50%), a portion of the overwintered cohort developed into adults the following spring; percentage of survival was negatively correlated to daily minimum temperature. P. humilis and P. lounsburyi successfully oviposited into host larvae in late fall, and their progeny developed into adults the following spring, although with a low percentage (0-11.9%) survivorship. Overwintering survival of puparia of the olive fruit fly and immature larvae of P. humilis and P. lounsburyi (inside host puparia), buried in the soil, were tested at an interior valley and coastal site. Survival of olive fruit fly ranged from 0 to 60% and was affected by the trial date and soil moisture. Overwintering survival of both the fruit fly and tested parasitoids was lower at the colder interior valley than the coastal site; P. humilis immature stages had the highest mortality levels while B. oleae pupae had the lowest mortality levels. The spring emergence pattern of the tested insects was well predicted by a degree-day model. We discuss factors potentially impeding establishment of introduced olive fruit fly parasitoids in California and elsewhere.
A two-year study was conducted in a citrus orchard, Citrus sinensis L., to determine frequency of predation on glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar). A total of 1,578 arthropod predators, representing 18 taxa, were collected and assayed for the presence of GWSS egg protein by an enzyme-linked immunosorbent assay using a Homalodisca -species and egg-specific monoclonal antibody and then by polymerase chain reaction using a H. vitripennis -specific DNA marker. The gut content analyses revealed the presence of GWSS remains in the gut of 2.28 % of the total arthropod predator population, with 3.09 % of the spiders and 0.59 % of the insect predators testing positive. Moreover, a comparison of the two assays indicated that they were not equally effective at detecting GWSS remains in predator guts. Low frequencies of GWSS detection in the gut of predators indicated that GWSS are not a primary prey and that predators may contribute little to suppression of this pest in citrus.
This study compares the fundamental spray characteristics, such as penetration depth, droplet diameter, velocity, and volume flux of, biodiesel and diesel as well as their blends. Liquid jet is injected into an air crossflow with a speed of 58 m/s. A series of experimental tests covering Weber numbers from 29-82, and momentum flux ratios from 10-133, are conducted using the shadowgraph method and phase Doppler particle analyzer. The shadowgraph images are processed to find the penetration depth and to investigate different regimes of breakup. The phase Doppler particle analyzer is used to capture droplet size distribution, velocity, and volume flux. The results differ remarkably in terms of breakup regimes and trajectory for biodiesel blends, which may be due to the high viscosity of biodiesel in comparison to diesel. For example, at a Weber number of 80, some bag breakups are observed on the biodiesel jet against the expectation that, at this Weber number, most of the liquids experience atomization mode.
Effects of cold storage temperatures and storage duration were evaluated for Psyttalia humilis (Silvestri) from Namibia and Psyttalia ponerophaga (Silvestri) from Pakistan, braconid parasitoids of Bactrocera oleae (Rossi) imported to California, USA. Immature stages of P. humilis were exposed to 4, 6, 8, 10, or 12 °C for 1, 2 or 4 months (pupa only at 4 and 12 °C) and then held at 24 °C for adult emergence. Less than 5 % of parasitoids in the 4–8 °C treatments survived, regardless of storage duration. At the 10 °C treatment, adult survival decreased with increased storage duration, but increased with advancing developmental stages. Survival was not affected at the 12 °C treatment. Adult P. humilis were exposed to 6, 8, 10 °C for short periods (1, 2, 4, or 6 weeks) or ambient winter conditions in Parlier, California, USA (about 9 °C). Regardless of storage temperature, P. humilis reproduction was reduced after storage of four and six weeks. Similarly, after 4 months at ambient winter temperatures, P. humilis reproduction was reduced. Psyttalia ponerophaga pupae stored at 6 °C for 41–97 days had decreased survival and increased developmental time. Survival of P. ponerophaga pupae ranged from 13.9–52.1 %, whereas under similar storage conditions survival of P. humilis was <0.7 %, suggesting P. ponerophaga is more cold tolerant than P. humilis.
Effects of ambient spring air temperature and light intensity on stylet penetration activities of the glassy-winged sharpshooter, Homalodisca vitripennis (Germar), were studied under field conditions by using an electrical penetration graph. Electrical penetration graph waveforms representing salivary sheath formation and searching (pathway phase), xylem contact (X waves), and ingestion of xylem fluid (waveform C) were analyzed. Previous research supported the concept that acquisition of Xylella fastidiosa, the Pierce's disease bacterium, occurs during ingestion, whereas inoculation occurs during xylem contact periods (X waves). Diel patterns of H. vitripennis stylet activity showed that, regardless of light condition, xylem ingestion occurred for the longest duration when temperature remained above the feeding threshold (10°C), and only occurred at temperatures below the threshold when ingestion was continued from a preceding, warmer time. Regression analysis indicated that mean waveform durations per insect (WDI) for combined stylet activities (pathway and ingestion) as well as X wave frequencies were significantly influenced by temperature, but there was no significant impact of light intensity or interaction between temperature and light intensity. The relationship between temperature and stylet activities in terms of WDI and X wave frequency was described using linear and nonlinear models. Validation of the nonlinear models indicated that they well predicted the WDIs for both ingestion and combined stylet activities, using temperature only as a single input. Overall, findings clearly demonstrate that temperature is an important factor that influences the H. vitripennis feeding behaviors responsible for transmission (acquisition and inoculation) of the Pierce's disease bacterium, with implications for vector ecology and management, as well as disease epidemiology.
The effects of temperature on the development (egg–adult emergence) of Gonatocerus morgani Triapitsyn, a newly-described parasitoid of Homalodisca vitripennis (Germar), were determined at 14.8, 18.7, 23.5, 26.9, 28.7, 30.4, 32.8, and 33.8°C in the laboratory. Survival rate (percent adult emergence from parasitized host eggs) varied significantly among the experimental temperatures, with the highest (59%) and lowest (0%) occurring at 30.4 and 33.8°C, respectively. The survival rates (%) were fitted with a polynomial model to describe a temperature-dependent pattern. Developmental rates (1/d) across seven temperatures were fitted with the nonlinear Briere model, which estimated the lower threshold to be 8.06°C, the optimal temperature to be 29.22°C, and the upper threshold to be 33.49°C. A linear model fitted to developmental rates at 14.8–28.7°C indicated that 189.75degree-days above the lower threshold of 9.71°C were required to complete development. A simulation model of G. morgani adult emergence was constructed to predict daily counts over the entire range of constant temperatures by incorporating the survival rate model, the Briere model, and the Weibull model. In outdoor validation, a degree-day model for predicting adult emergence showed ⩽2d differences between prediction and observation. Based on the observed temperature requirement, the insect could complete thirteen to sixteen generations per year in southern California, depending on weather and location.
The parasitoid Psyttalia humilis (Silvestri) was reared on Mediterranean fruit fly, Ceratitis capitata (Wiedemann), third instars irradiated at 0-70 Gy at the USDA, APHIS, PPQ, Moscamed biological control laboratory in San Miguel Petapa, Guatemala, and shipped to the USDA, ARS, Parlier, CA. Irradiation dose did not affect the parasitoid's offspring sex ratio (53-62% females), percentage of unemerged adults (12-34%), number of progeny produced per female (1.4-1.8), and parasitism (19-24%). Host irradiation dose had no significant effect on the forewing length of female P. humilis and its parasitism on olive fruit fly, Bactrocera oleae (Rossi) and offspring sex ratio, but dissection of 1-wk-old female parasitoids reared from hosts irradiated with 70 Gy had a significantly lower number of mature eggs than females from nonirradiated hosts. Longevity of P. humilis adults decreased with increased temperature from 15 to 35°C, regardless of food provisions, gender, and host irradiation dose. Females survived 37-49 d at 15°C with water and food, and only 1-2 d at 35°C without food, whereas males lived shorter than females at all temperatures and food combinations tested. Adult P. humilis reared from fertile C. capitata and aspirated for dispensing in cups lived significantly longer after shipment than those specimens chilled and dispensed by weight. At 21 and 32°C, 50% of parasitoids departed release cages after 180 and 30 min, respectively, but none departed at 12°C. Thirteen shipments of P. humilis (2,980-21,922 parasitoids per shipment) were received between September and December 2009, and seven shipments (7,502-22,560 parasitoids per shipment) were received between October and December 2010 from San Miguel Petapa, Guatemala. Daily number of olive fruit fly adult and percentage female trap captures ranged <1-19 and 8-58% in 2009, and <1-11 and 0-42% in 2010, respectively. The number of parasitoids released ranged 848-12,257 in 2009 and 3,675-11,154 in 2010. Percentage parasitism of olive fruit fly third instars at all locations ranged 0-9% in 2009 and 0-36% in 2010.
ABSTRACT Homalodisca vitripennis (Germar), a vector of Xylella fastidiosa, is associated with citrus plantings in California. Infested citrus orchards act as a source of vectors to adjacent vineyards where X. fastidiosa causes Pierce's disease. An analysis of the pattern and rate of movement of H. vitripennis and its egg parasitoid, Gonatocerus ashmeadi Girault, was conducted in a citrus orchard by using a protein mark-capture technique to quantify movement and net dispersal rates in the experimental areas. Treatments included irrigation at 100% of the crop evapotranspiration rate (ETc), 80, and 60% ETc. Sex-specific net dispersal rates showed that H. vitripennis males and females moved consistently and contributed equally to the level of population change within treated areas. Trees irrigated at 60% ETc were the least preferred by H. vitripennis. Among all protein-marked individuals captured in the 60% ETc treatment, ≈75 and 88% in 2005 and 2006, respectively, were inflow individuals. Movement toward less preferable plants indicates that in agricultural landscapes dominated by perennial monocultures, there is a random component to H. vitripennis movement, which may result from the inability of H. vitripennis to use plant visual cues, olfactory cues, or both to make well-informed long-range decisions. The 80% ETc areas were a significant source of adult H. vitripennis and G. ashmeadi compared with the other treatments. Colonization rates by parasitoids were synchronized with the spatiotemporal distribution of H. vitripennis eggs. Results suggest that H. vitripennis movement from citrus into adjacent vineyards could be a result of random dispersal rather than oriented movement in response to host-plant characteristics.
Long-term separation of a host from its native parasitoids may result in divergent thermal adaptation between host and parasitoid. The olive fruit fly, Bactrocera oleae (Rossi), most likely originated from Sub-Saharan Africa, but has since had a long invasion history in cultivated olives that spans geographical barriers and continents. This study compared three major thermal performance profiles (development, survival, and reproduction) across a wide range of temperatures (10–34 °C) among a Californian population of the olive fruit fly and two African parasitoids, Psyttalia lounsburyi (Silvestri) and Psyttalia humilis (Silvestri), believed to have co-adapted with the fruit fly in its native range. Temperature ranges for the development and survival were 10–30 °C for the fly, 10–28 °C for P. lounsburyi, and 14–32 °C for P. humilis. There was no difference in any thermal performance measured between two P. humilis populations (Kenya and Namibia) tested. The most suitable temperature ranges for reproduction were 22–30 °C for the fly, 18–32 °C for P. humilis, and 18–26 °C for P. lounsburyi. The results showed slight differences in the thermal profiles among olive fruit fly and both parasitoids species, with P. humilis being more heat tolerant whereas P. lounsburyi was less heat tolerant than the fruit fly. The results are discussed with respect to thermal co-adaptation and classical biological control of the olive fruit fly.
The larval parasitoid, Psyttalia humilis (Silvestri), reared on Mediterranean fruit fly, Ceratitis capitata (Weidemann), by USDA-APHIS-PPQ, San Miguel Petapa, Guatemala was imported into California for biological control of olive fruit fly, Bactrocera oleae (Rossi). This study reports the results of field releases and recovery of P. humilis in California, and laboratory investigations to determine the effects of food provision, high temperature, and insecticidal bait spray on the parasitoid's survival and fecundity. Parasitoids (3,613-7,823) were released in Orland, San Juan Bautista, Cayucos, Sylmar, Santa Barbara, and San Diego during October through December 2006. Mean daily temperatures at the release sites ranged from 10.7 degrees C in Orland to 20.9 degrees C in San Juan Bautista. The lowest (0.5) and highest (29.7) mean number of adult B. oleae per day per trap was captured in Orland and Sylmar, while the lowest (0.01) and highest (2.21) number of third instar larvae per fruit was collected on 11 December in Orland and on 5 October in San Diego in prerelease samples, respectively. Parasitoids were recovered from all release sites, the lowest (0.3%) and highest (100%) parasitism occurred on 25 January in Sylmar and on 26 October in Cayucos, respectively. At 24 degrees C, parasitoids reared from B. oleae larvae survived 36 d on honey, 31 d on orange juice, and 28 d on honeydew, which was significantly longer than on cut olive fruit (8 d) or without food (11 d), but was similar to those reared from C. capitata larvae under the same food conditions. Under a high diurnal temperature regime (18.3-35 degrees C) reflecting the summer olive growing conditions in the California Central Valley, the parasitoids survived <5 d when no food or only water was provided. Its longevity and life-time fecundity significantly increased by provision of honey or honeydew. There was no difference in the parasitoid's longevity between females and males or between food and sexes. Percent mortality of parasitoid adults was not significantly affected by the exposure to insecticidal fruit fly bait (GF-120) in four different types of choice tests with artificial honeydew and GF-120.