Lincoln Smith

Influence of Seed Head–Attacking Biological Control Agents on Spotted Knapweed Reproductive Potential in Western Montana Over a 30-Year Period

Abstract Five insect biological control agents that attack flower heads of spotted knapweed, Centaurea stoebe L. subsp. micranthos (Gugler) Hayek, became established in western Montana between 1973 and 1992. In a controlled field experiment in 2006, seed-head insects reduced spotted knapweed seed production per seed head by 84.4%. The seed production at two sites in western Montana where these biological control agents were well established was 91.6–93.8% lower in 2004–2005 than 1974–1975, whereas the number of seed heads per square meter was 70.7% lower, and the reproductive potential (seeds/m2) was 95.9–99.0% lower. The average seed bank in 2005 at four sites containing robust spotted knapweed populations was 281 seeds/m2 compared with 19 seeds/m2 at four sites where knapweed density has declined. Seed bank densities were much higher at sites in central Montana (4,218 seeds/m2), where the insects have been established for a shorter period. Urophora affinis Frauenfeld was the most abundant species at eight study sites, infesting 66.7% of the seed heads, followed by a 47.3% infestation by Larinus minutus Gyllenhal and L. obtusus Gyllenhal. From 1974 to 1985, Urophora spp. apparently reduced the number of seeds per seed head by 34.5–46.9%; the addition of Larinus spp. further reduced seed numbers 84.2–90.5% by 2005. Path analysis indicated that both Larinus spp. and U. affinis contributed significantly to reduction of seed production over the 30-yr period. Spotted knapweed density may not decrease significantly until the seed bank falls below a critical threshold.

In situ volatile collection, analysis, and comparison of three Centaurea species and their relationship to biocontrol with herbivorous insects.

Centaurea solstitialis, commonly known as yellow starthistle, is an invasive plant listed as a noxious weed in the western areas of North America and is the target of classical biological control, which involves release of herbivores known to be specific to this plant. These insects often choose their host plant on the basis of the volatile organic compounds (VOCs) emitted. Accordingly, volatile analysis of host plants can provide insight into VOCs that may attract and/or repel the insect. To this end, solid-phase microextraction (SPME) and a customized collection bag were utilized to perform in situ volatile collection on intact and mechanically damaged leaves of Centaurea solstitialis, Centaurea cyanus, and Centaurea cineraria. Volatile identification was performed by GC-MS, and the VOC differences were determined. The plants C. solstitialis and C. cyanus have been reported to attract the weevil, Ceratapion basicorne, a candidate for biological control, whereas C. cineraria does not attract the weevil. Major VOCs unique to C. cineraria include the sesquiterpenes cyclosativene, alpha-ylangene, and trans-alpha-bergamotene. The compound trans-beta-farnesene was unique to C. solstitialis and C. cyanus.

Field cage assessment of interference among insects attacking seed heads of spotted and diffuse knapweed

Abstract Field studies were conducted to determine the competitive interactions between introduced biological control agents that attack the seed heads of spotted knapweed (Centaurea stoebe ssp. micranthos) and diffuse knapweed (Centaurea diffusa). Two weevils, Bangasternus fausti and Larinus minutus (Coleoptera: Curculionidae), were each paired with the previously established fly, Urophora affinis (Diptera: Tephritidae). Each species was released either alone or in pair-wise combinations inside screen cages placed over existing knapweed plants at six field sites in Montana and one in Oregon. Larinus minutus produced almost three times as many progeny on diffuse knapweed as on spotted knapweed. Larinus minutus reproduction was not affected by competition with U. affinis, but U. affinis reproduction was reduced by the presence of L. minutus (by 71% on spotted and 77% on diffuse knapweed). Bangasternus fausti reproduction generally was not affected by competition with U. affinis, nor was U. affinis affected by B. fausti on either host plant. There were extremely few cases of successful production of both weevil and fly in the same capitulum, which was probably because weevil larvae consume the developing flies. Both weevils increased the total proportion of seed heads infested on diffuse knapweed, and B. fausti increased it on spotted knapweed. However, the release of either weevil did not significantly further reduce seed production on either plant. The results and experimental design are discussed in light of the subsequent establishment and impact of these agents.

Population densities of yellow starthistle (Centaurea solstitialis) in Turkey

Abstract Yellow starthistle is one of the most important alien invasive weeds in the western United States. It has been targeted for biological control based on the assumption that its abundance is limited by natural enemies in its native region but not in the United States. The geographic center of diversity for yellow starthistle appears to be in Turkey. This region is being explored to discover potential biological control agents; however, there is no quantitative information regarding the population density or dynamics of the plant in this region. Such information could help determine which natural enemies help suppress the plant in its land of origin. We measured densities of yellow starthistle plants and seeds during 2 yr at three locations in central Turkey. Densities of mature plants were about 4% of those measured at sites in California. Densities of capitula and seeds produced were about 60 and 65%, respectively, of those measured in California. The greatest difference between the two regions appears to be the densities of mature plants, which indicates the importance of focusing research on natural enemies that reduce plant survival. Nomenclature: Yellow starthistle, Centaurea solstitialis L. CENSO.

Relationship Among Growth Attributes of Spotted Knapweed (Centaurea maculosa) in Western Montana1

Spotted knapweed is an important weed of rangeland in the northwestern United States. A study was conducted near Corvallis, MT, during 1992 to 1994 in order to assess the relationship among the growth attributes of spotted knapweed to identify a minimum set of measurable plant characteristics that are representative of spotted knapweed vigor. Spotted knapweed growth attributes that were examined included plant age, root diameter, plant height, number of stems per plant, aboveground biomass, number of capitula (seed heads) per plant, and number of capitula per stem. Spotted knapweed age was positively correlated with root diameter, number of stems per plant, aboveground biomass, and proportion of bolted plants. Most spotted knapweed plants did not bolt until the third or fourth year. Although plant age is not measured easily in the field, it may be useful as a covariate in an analysis of experiments involving plant competition or nonlethal biological control agents. Root diameter can be used as a nondestructive measure of approximate plant age, especially for the first 5 yr of growth. Root diameter was also highly correlated with many growth measurements, including number of capitula per plant and aboveground biomass, which are most relevant to assessing overall plant vigor. Plant height was positively correlated with aboveground biomass, number of capitula per plant, and mean number of capitula per stem. Number of stems per plant was positively correlated with plant height, aboveground biomass, and number of capitula per plant. Aboveground biomass was positively correlated to number of capitula per plant and mean number of capitula per stem. Measurements of root diameter, plant height, and number of stems are easy to perform and should provide a good indication of plant vigor. Nomenclature: Spotted knapweed, Centaurea maculosa Lam. #3 CENMA. Additional index words: ANCOVA, analysis of covariance; growth–attribute relationships.

Effect of mechanical damage on emission of volatile organic compounds from plant leaves and implications for evaluation of host plant specificity of prospective biological control agents of weeds

Abstract Assessment of host plant specificity is a critical step in the evaluation of classical biological control agents of weeds which is necessary for avoiding possible damage to non-target plants. Volatile organic compounds (VOCs) emitted by plants likely play an important role in determining which plants attract and are accepted by a prospective arthropod agent. However, current methods to evaluate host plant specificity usually rely on empirical choice and no-choice behavioural experiments, with little knowledge about what chemical or physical attributes are stimulating the insect. We conducted experiments to measure the quantitative and qualitative effects on emission of VOCs caused by simple mechanical damage to leaves of plants known to differ in suitability and attractiveness to a prospective agent. More VOCs were detected from damaged than from undamaged leaves for all three species tested. Discriminant analysis was able to correctly distinguish the taxonomic identity of all plants based on their VOC profiles; however, the VOCs that discriminated species among undamaged leaves were completely different from those that discriminated among damaged leaves. Thus, damaged and undamaged plants present different VOC profiles to insects, which should be considered when conducting host plant specificity experiments. An unacceptable non-target plant, Centaurea cineraria, emitted all except one of the VOCs that were emitted by its preferred host plant, Centaurea solstitialis, indicating the importance of compounds that are repellant in host plant specificity. Centaurea cyanus emitted fewer VOCs than C. solstitialis, which suggests that it lacked some VOCs important for host plant recognition.

Suitability of the maize weevil and angoumois grain moth as hosts for the parasitoids Anisopteromalus calandrae and Pteromalus cerealellae

Two parasitoids, Pteromalus cerealellae (Ashmead) and Anisopteromalus calandrae (Howard) (Hymenoptera: Pteromalidae), were compared for their ability to parasitize two important internally‐developing insect pests of stored maize (Zea mays L.). Parasitism by P. cerealellae was greater on Angoumois grain moth, Sitotroga cerealella (Olivier), than on maize weevil, Sitophilus zeamais Motschulsky, in no‐choice experiments. Anisopteromalus calandrae parasitized more maize weevils than did P. cerealellae. The former parasitoid parasitized only a few Angoumois grain moths successfully in maize, but parasitized many in wheat if the hosts were younger than 3 weeks old. Thus, both host age and type of grain affect suitability for parasitism. The effects of parental host (species on which the female developed) and experimental host (species exposed to parasitism) on parasitism rate of P. cerealellae were tested in a host‐switching experiment. Parasitism by parasitoids reared on maize weevils was 23% lower than that of parasitoids reared on Angoumois grain moth. This effect was independent of which host the filial generation of parasitoids was tested on. However, the experimental host species had a much greater effect on parasitoid fecundity than the parental host species. Female progeny had smaller body sizes when emerging from maize weevil than from Angoumois grain moth, which may explain the parental host effect on fecundity. There was also a slight intergenerational effect of host species on parasitoid body size.

Fecundity, Development, and Behavior of Ceratapion basicorne (Coleoptera: Apionidae), a Prospective Biological Control Agent of Yellow Starthistle

Abstract A laboratory colony of Ceratapion basicorne (Illiger) was established from adults reared from infested plants of yellow starthistle, Centaurea solstitialis L. (Asteraceae), that were collected in eastern Turkey. Newly emerged adults fed on yellow starthistle foliage and mated, but females did not oviposit. The feeding rate of females decreased to almost zero by 16 d after emergence, whereas males continued to feed for at least 26 d. Dispersal activity was initially high but decreased to low levels in 2–3 wk. After 6 wk, most adults were hiding inside tightly curled dry leaves and in the crevices of crumpled paper towel. Insects were held in a cold dark incubator (5°C) for at least 3 mo to terminate reproductive diapause. Females began ovipositing 4.4 d after being placed on yellow starthistle leaves at 19°C. The oviposition period lasted a mean of 20.6 d, and lifetime fecundity was 34.5 eggs. Daily fecundity was high during the first 14 d of oviposition (1.8 eggs/d) and declined to low levels (0.3 eggs/d). Feeding rate of ovipositing females during the first 2 wk was 19.2 holes/d, but this decreased to 4.7 holes/d for the remainder of the 45-d experiment. Female feeding rate was highly correlated to oviposition rate. Development time of eggs until eclosion of larvae at 19°C was 8.5 d and survivorship until eclosion was 73%. Development time from oviposition until adult emergence at ≈19°C was 77 d. These results provide a foundation for conducting experiments to evaluate host plant specificity and potential impact on the weed.

Field assessment in land of origin of host specificity, infestation rate and impact of Ceratapion basicorne a prospective biological control agent of yellow starthistle

Yellow starthistle, Centaurea solstitialis (Asteraceae), is an important invasive alien weed in the western United States. Currently established biological control agents attack only the capitula (flowerheads), and are not effectively controlling the plant in much of its range. The geographic center of diversity for the plant appears to be in Turkey, but no agents have been introduced from this country. Ceratapion basicorne (Coleoptera: Apionidae) is common in Central Turkey, attacking 25–100% of yellow starthistle plants. In a field experiment, Ceratapion spp. attacked 90% of yellow starthistle plants and 88% of milk thistle plants (Silybum marianum) but not seven other plant species, including artichoke and safflower. We suspect that a different species of insect attacked milk thistle, but they emerged before the plants were sampled. Laboratory tests showed that C. basicorne does not oviposit in milk thistle. Ceratapion basicorne appears to be more host specific than was suggested by previous studies of a population in Italy (Clement et al. 1989. Ann. Entomol. Soc. Am. 82: 741–747). The insect is gregarious, and the number of larvae per plant was positively correlated to root diameter. The level of damage to individual plants was positively correlated to the proportion of plants attacked, indicating aggregation both among plants and within plants. Field data did not show any impact of the insect on plant size or number of capitula, but germination rate of seeds produced by infested plants was 15% lower than for uninfested plants at two of three sites studied.

Host plant oviposition preference of Ceratapion basicorne (Coleoptera: Apionidae), a potential biological control agent of yellow starthistle.

Abstract Ceratapion basicorne is a weevil native to Europe and western Asia that is being evaluated as a prospective classical biological control agent of Centaurea solstitialis (yellow starthistle) in the United States. Choice oviposition experiments were conducted under laboratory conditions to help assess host-plant specificity of the insect. Mean oviposition rates were highest on C. solstitialis (66% of eggs, on a per replicate basis) followed by Centaurea cyanus (bachelors button 22%), Centaurea melitensis (6%), Centaurea americana (1%), Saussurea americana (3%) and Carthamus tinctorius (safflower 2%). Adult feeding damage followed a similar pattern; however, there was less oviposition relative to the amount of adult feeding on each of the non-target species than on the target host plant, C. solstitialis. Thirteen safflower varieties were tested, and oviposition occurred on eight of them, at low rates. Adult feeding occurred on all safflower varieties tested, although at rates much lower than on yellow starthistle. The results were intermediate between those of previously reported no-choice laboratory and open field experiments. Overall, the combined results support the hypothesis that C. basicorne is not likely to attack any of the non-target plant species tested here except possibly C. cyanus and C. melitensis, which are both invasive alien plants.