Influence of Centaurea biebersteinii patch size on Urophora quadrifasciata (Dipt. Tephritidae) in Michigan, USA

Abstract:  Host plant patch size plays an important role in the distribution and densities of many insect species. Understanding how the host plant patch size influences the population of a biological control agent is necessary to monitor the success of a biological control programme. Urophora quadrifasciata (Dipt. Tephritidae) was released in the early 1970s to control the spotted knapweed, Centaurea biebersteinii (Asteraceae) in North America. The studies reported here investigated the influences of spotted knapweed patch size and stem density on U. quadrifasciata larval populations overwintering within capitula in the Upper Peninsula of Michigan, USA. An inverse relationship was found between the percentage of capitula with U. quadrifasciata larvae and spotted knapweed patch size in 2002 from Houghton county, MI. In 2003, an inverse relationship was also found in the south region of the study area in Houghton and Keweenaw counties, Michigan. The percentage of capitula with U. quadrifasciata larvae and spotted knapweed patch size were positively related for the central region. No relationship was found for the North region in 2003. Dispersal of U. quadrifasciata is directed north-east along the Keweenaw Peninsula, Michigan. With this directed spread, areas with well-established populations of U. quadrifasciata would provide source individuals for new colonization. Spotted knapweed patch size may play an important role in the overwintering populations of U. quadrifasciata as a result of dispersal.     

Influence of seed head-attacking biological control agents on spotted knapweed reproductive potential in western Montana over a 30-year period

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/m(2)) was 95.9-99.0% lower. The average seed bank in 2005 at four sites containing robust spotted knapweed populations was 281 seeds/m(2) compared with 19 seeds/m(2) at four sites where knapweed density has declined. Seed bank densities were much higher at sites in central Montana (4,218 seeds/m(2)), 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.     

Establishment and Dispersal ofUrophora affinis(Diptera: Tephritidae) andMetzneria paucipunctella(Lepidoptera: Gelechiidae) in Southwestern Virginia

Two European gall-producing insects,Urophora affinisFrfld. (Diptera: Tephritidae) andMetzneria paucipunctella(Zeller) (Lepidoptera: Gelechiidae) were introduced into Virginia in 1986 for biological control of spotted knapweed (Centaurea maculosaLam.). Adults ofU. affinis(n = 2625) andM. paucipunctella(n = 450) were released at two sites in Montgomery County, Virginia, and their populations were monitored yearly by dissecting spotted knapweed flower heads. Beginning in 1992, knapweed samples collected at various distances from the release sites were checked for dispersal.U. affinisis well established and is spreading slowly. The number of larvae per flower head and the seed numbers are inversely related as plants with the greatest number of larvae per spotted knapweed head had the lowest number of seeds. Knapweed density has declined at one of the release sites which had the highest rate of infestation byU. affinis.Establishment of the moth,M. paucipunctella,is less certain as it has been recovered at a very low level from only one site.     

Host plant testing ofUrophora quadrifasciata [Diptera: Tephritidae] againstCarthamus tinctorius and two North American species ofCentaurea

Host specificity ofUrophora quadrifasciata (Meig.) [Dipt.: Tephritidae], an introduced biological control agent of diffuse and spotted knapweed, was investigated by exposing flies to safflower,Carthamus tinctorius L., and 2 native knapweeds,Centaurea americana Nutt. andC. rothrockii Greenman. Results of the tests, and literature records indicate thatU. quadrifasciata is host specific to a few closely related plants of introduced knapweed species, and thus poses no threat to economically and ecologically important plant species in the United States.      

Role of plant phenology in mediating interactions between two biological control agents for spotted knapweed

The role of spotted knapweed phenology on the attack rate of two seed-head insects Urophora affinis and Larinus minutus was assessed in a series of field studies at four study sites in south-eastern British Columbia, Canada. Slow or later developing knapweed plants had more seed heads that contained only single or multiple U. affinis whereas early or faster developing plants had more seed heads containing L. minutus alone or in combination with U. affinis. L. minutus did not distinguish between seed heads with or without U. affinis larvae when laying eggs. However, seed heads with multiple U. affinis present, produced fewer L. minutus adults than expected. The probability of single or multiple U. affinis galls being present increased with seed-head diameter but was not affected by seed-head height. Attack by L. minutus increased with seed-head diameters >5 mm and was lower at plant heights above 50 cm. These results demonstrate two mechanisms that enable U. affinis to successfully coexist with L. minutus: differences between the species in their response to the developmental phenology of knapweed heads, and increased survivorship of U. affinis in heads with multiple U. affinis galls through niche interference competition. These mechanisms provide a possible explanation for the persistence of U. affinis populations on spotted knapweed, in spite of high levels of within seed-head mortality that have been observed with increasing L. minutus populations.     

A revision of the western Palaearctic species of Urophora Robineau-Desvoidy (Diptera: Tephritidae)

Abstract A key is provided to twenty-four western Palaearctic species of Urophora Robineau-Desvoidy. The hosts of twenty-three species which attack Asteraceae are listed, including those being used or investigated as possible weed biocontrol agents. The species are divided into four species groups and the differing host relationships and types of galls induced by these groups are discussed. U.lopholomae sp.n. and U.affinis ssp. calcitrapae ssp.n., associated with Centaurea (Lopholoma) spp. and C. (Calcitrapa) spp. respectively, are described. U. algerica (Hering) and U.sjumorum (Rohdendorf) are both treated as subspecies of U. quadrifasciata (Meigen). U.pontica is given full specific status and U.hispanica is removed from synonymy. The following new synonymies are made (junior synonyms in parentheses): U. angustifascia (Hering) (= Euribia phaeocera Hering); U. cardui (Linnaeus) (= U. reaumurii Robineau-Desvoidy, lectotype designated); U. jaceana (Hering) (= E.conyzae Hering); U.maura (Frauenfeld) (= E. tecta Hering); U. mauritanica Macquart (= U. lejura Rondani, Trypeta macrura Loew); U.solstitialis (Linnaeus) (= E.sonderupi Hering, U. veruata Rondani ); U.stylata (Fabricius) (= E.pia Hering, U. vulcaanica Rondani); U. terebrans (Loew) (= E. approximata Hering, T. eriolepidis Loew, E. manni Hendel). The possibility that U. quadrifasciata is a species complex is discussed; it is also suggested that U.affinis and U.jaceana represent the morphological extremes of a complex. The misuse of the name Musca stylata Fabricius in the genus Myopites Blot is noted.     

Non-target effects of an introduced biological control agent on deer mouse ecology

Release of exotic insects as biological control agents is a common approach to controlling exotic plants. Though controversy has ensued regarding the deleterious direct effects of biological control agents to non-target species, few have examined the indirect effects of a ”well-behaved” biological control agent on native fauna. We studied a grassland in west-central Montana infested with spotted knapweed (Centaurea maculosa) to examine the effects of knapweed invasion and two gall flybiological control agents (Urophora affinis and U. quadrifasciata) on the native deer mouse (Peromyscus maniculatus). Stomach-content analysis revealed that Urophora were the primary food item in Peromyscus diets for most of the year and made up 84–86% of the winter diet. Stomach contents indicated that wild-caught mice consumed on average up to 247 Urophora larvae mouse^–1 day^–1, while feeding trials revealed that deer mice could depredate nearly 5 times as many larvae under laboratory conditions. In feeding trials, deer mice selected knapweed seedheads with greater numbers of galls while avoiding uninfested seedheads. When Urophora larvae were present in knapweed seedheads, deer mice selected microhabitats with moderately high (31–45% cover) and high knapweed infestation (≥46% cover). After Urophora emerged and larvae were unavailable to Peromyscus, mice reversed habitat selection to favor sites dominated by native-prairie with low knapweed infestation (0–15%). Establishment of the biological control agent, Urophora spp., has altered deer mouse diets and habitat selection by effecting changes in foraging strategies. Deer mice and other predators may reduce Urophora populations below a threshold necessary to effectively control spotted knapweed. Received: 04 May 1999 / Accepted: 14 August 1999     

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

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.     

Arbuscular mycorrhizal fungi enhance spotted knapweed growth across a riparian chronosequence

Arbuscular mycorrhizal fungi (AMF) mediate nutrient uptake that accelerates plant growth and reproduction. Thus, AMF may promote plant invasions often observed along rivers. We assessed the importance of AMF in improving growth of the invasive species, spotted knapweed (Centaurea stoebe), during succession of riparian vegetation along a flood plain in Montana, USA. We grew spotted knapweed with and without AMF in soils collected from riparian sites ranging from 1 to 72 years old and measured the plant’s growth response to AMF. We observed variability in relative effects of AMF, with greatest growth benefits in recently deposited alluvial sediments. We then separated effects of soil and inoculum source by growing spotted knapweed with soils and inocula collected from young or old sites and found that growth responses were greatest in young soils regardless of inoculum source. Our results demonstrate that AMF directly benefit growth of spotted knapweed, especially in soils that typify early successional sites on this alluvial flood plain.     

Growth and development of the knapweed root weevil,Cyphocleonus achates,on a meridic larval diet

Cyphocleonus achates, the knapweed weevil, is an effective biological control agent of the invasive weed, Centaurea maculosa Lam. A meridic diet was developed and tested for the rearing of the larval stage of this insect. Using this diet, C. achates was reared for over three generations, with the adults being offered knapweed plants for feeding and oviposition in greenhouse conditions. Slight or no differences were seen between insects reared on a standard meridic diet formulation and one containing knapweed tissues. The following life history parameters were monitored over the three generations: percent egg hatch (ranging from 42.9 to 59.1%), time to egg hatch (20.0–23.2 days), time to adult emergence (52.0–54.1 days), adult weights 3 days post-eclosion (101.9–117.0 mg), percent adult emergence (48.3–58.6%), and percent mortality/deformity in the different stages (with mortality occurring primarily in the early larval stages). Additionally, a study involving low temperature and short day conditions suggested that C. achates could be maintained for longer periods of time in larval diet cells when placed in growth-retarding conditions, although percent adult emergence was lower. External morphology was also studied in order to distinguish between the sexes to ensure that each adult cage had a similar ratio of females to males. Abdominal features were found to be the most dependable characteristics for use when determining the sex of adult C. achates.     

How do biological control and hybridization affect enemy escape?

Two mechanisms often linked with plant invasions are escape from enemies and hybridization. Classical biological control aims to reverse enemy escape and impose top-down population control. However, hybridization has the potential to alter interactions with enemies and thus affect biological control. We examined how introductions of biological control agents affect enemy escape by comparing specialist enemy loads between the native and introduced ranges of two noxious weeds (spotted and diffuse knapweed; Centaurea stoebe L. and C. diffusa Lam.) that have been the targets of an extensive biological control program. Hybrids between spotted and diffuse knapweed are often found within diffuse knapweed sites in North America, so we also compared enemy loads on plants that appeared morphologically like diffuse knapweed and hybrids. Finally, we tested the preference for diffuse knapweed, hybrids, and spotted knapweed of one of the agents thought to be instrumental in control of diffuse knapweed (Larinus minutus; Curculionidae). In North America spotted knapweed has largely escaped its root herbivores, while seedhead herbivore loads are comparable in the introduced and native ranges. Diffuse knapweed exhibited seedhead herbivore loads five times higher in the introduced compared to native range. While this pattern of seedhead herbivory is expected with successful biological control, increased loads of specialist insect herbivores in the introduced range have rarely been reported in the literature. This finding may partially explain the better population control of diffuse vs. spotted knapweed. Within North American diffuse knapweed sites, typical diffuse knapweed and hybrid plants carried similar herbivore loads. However, in paired feedings trials, the specialist L. minutus demonstrated a preference for newly created artificial hybrids over North American diffuse knapweed and for European diploid spotted knapweed over North American tetraploid spotted knapweed. Overall though, hybridization does not appear to disrupt biological control in this system.     

Biological control agents elevate hantavirus by subsidizing deer mouse populations

Biological control of exotic invasive plants using exotic insects is practiced under the assumption that biological control agents are safe if they do not directly attack non-target species. We tested this assumption by evaluating the potential for two host-specific biological control agents ( Urophora spp.), widely established in North America for spotted knapweed ( Centaurea maculosa ) control, to indirectly elevate Sin Nombre hantavirus by providing food subsidies to populations of deer mice ( Peromyscus maniculatus ), the primary reservoir for the virus. We show that seropositive deer mice (mice testing positive for hantavirus) were over three times more abundant in the presence of the biocontrol food subsidy. Elevating densities of seropositive mice may increase risk of hantavirus infection in humans and significantly alter hantavirus ecology. Host specificity alone does not ensure safe biological control. To minimize indirect risks to non-target species, biological control agents must suppress pest populations enough to reduce their own numbers.     

Interactions and effects of multiple biological control insects on diffuse and spotted knapweed in the Front Range of Colorado

Abundances and interactions among biological control insects and their effects on target invasive plants were monitored within the flower heads and roots of diffuse knapweed, Centaurea diffusa, and in spotted knapweed, Centaurea stoebe, along the Colorado Front Range. Flower weevils, (Larinus species) and root-feeders (Cyphocleonus achates and Sphenoptera jugoslavica) were released on knapweed that already supported biological control gall flies (Urophora species). At a single monitoring site, seed production by C. diffusa declined from 4400 seeds m⁻² in 1997 to zero seeds m⁻² on the monitoring sites in 2006, while the flowering stem density of C. diffusa declined from a peak of almost 30 stems m⁻² in 2000 to zero stems m⁻² in 2006. The average abundance of Urophora and Larinus in flower heads fluctuated independently during the 2001–2006 interval, while the relative abundance of C. achates and S. jugoslavica in roots exhibited a weak inverse relationship that appeared driven by climate effects. The relative abundance of insects on a population of C. stoebe was monitored for five years as Larinus species and C. achates became established on spotted knapweed that already supported Urophora species. Spotted knapweed seed production on our monitoring site declined from 4600 seeds m⁻² in 2003 to zero seeds m⁻² in 2006. Unlike C. diffusa, substantial numbers of rosettes of C. stoebe remained present. Larinus consumed almost all Urophora encountered in C. diffusa, and consumed about 40% of the Urophora in co-infested flower heads of C. stoebe (ca. 10–15% of the total Urophora population). No negative correlations between the relative densities of flower head and root-feeding insects were observed. The effects of these insects on target plants have produced results consistent with the ‘cumulative stress hypothesis’ for biological control of Centaurea species.     

Is spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis> L.) patch size related to the effect on soil and vegetation properties?

Spotted knapweed (Centaurea stoebe L. subsp. Micranthos (Gugler) Hayek) was first introduced in the 1890s from Europe into western North America, where it now occupies over three million hectares of rangeland and pasture in 14 states and two Canadian provinces, reducing forage production and causing economic damage. Despite many reported effects spotted knapweed can have on soils and native vegetation, it is not known whether patch size is correlated with these ecosystem-level effects. The objective of our study was to determine whether the effects of spotted knapweed on plant composition and soil properties was related to spotted knapweed patch size. We asked the following questions: (1) Are there differences in plant species richness and diversity between small and large knapweed patches? and (2) Do soil water and soil mineral nutrient properties change depending on knapweed patch size? Twenty-four knapweed patches, and paired natural grassland plots, were randomly selected within Lac du Bois Provincial Park, British Columbia, Canada. Knapweed patch size ranged from 6 to 366 m². Sampling and analysis revealed a significant effect of knapweed patch size on soil and vegetation properties. Soil P, soil temperature, and total dry plant biomass (g/0.25 m²) increased, while soil N, soil C, and soil moisture decreased with patch size. Since our results show that spotted knapweed patch size is related to degree of soil alteration, it is important to consider size of patch when modeling the impact of spotted knapweed in North America. Since large patches of spotted knapweed seem to have a proportionately greater effect on soil chemistry properties, large patches may move the system further away from a point where it is possible to restore the site to pre-invasion conditions.     

Invasion of an exotic forb impacts reproductive success and site fidelity of a migratory songbird

Although exotic plant invasions threaten natural systems worldwide, we know little about the specific ecological impacts of invaders, including the magnitude of effects and underlying mechanisms. Exotic plants are likely to impact higher trophic levels when they overrun native plant communities, affecting habitat quality for breeding songbirds by altering food availability and/or nest predation levels. We studied chipping sparrows (Spizella passerina) breeding in savannas that were either dominated by native vegetation or invaded by spotted knapweed (Centaurea maculosa), an exotic forb that substantially reduces diversity and abundance of native herbaceous plant species. Chipping sparrows primarily nest in trees but forage on the ground, consuming seeds and arthropods. We found that predation rates did not differ between nests at knapweed and native sites. However, initiation of first nests was delayed at knapweed versus native sites, an effect frequently associated with low food availability. Our seasonal fecundity model indicated that breeding delays could translate to diminished fecundity, including dramatic declines in the incidence of double brooding. Site fidelity of breeding adults was also substantially reduced in knapweed compared to native habitats, as measured by return rates and shifts in territory locations between years. Declines in reproductive success and site fidelity were greater for yearling versus older birds, and knapweed invasion appeared to exacerbate differences between age classes. In addition, grasshoppers, which represent an important prey resource, were substantially reduced in knapweed versus native habitats. Our results strongly suggest that knapweed invasion can impact chipping sparrow populations by reducing food availability. Food chain effects may be an important mechanism by which strong plant invaders impact songbirds and other consumers.     

Influence of Nutrient Availability on the Interaction Between Spotted Knapweed and Bluebunch Wheatgrass

Centaurea maculosa (Lam.) (spotted knapweed) reduces wildlife and livestock habitat biodiversity and increases erosion. Nutrient availability to plants may be used to accelerate succession away from spotted knapweed. Early‐successional plant communities often have high nutrient availability, whereas late‐successional communities are often found on lower nutrient soils. We hypothesized that removal of nutrients would change the competitive advantage from spotted knapweed to Pseudoroegneria spicatum (bluebunch wheatgrass) (late seral). In two addition series matrices, background densities of Secale cereale (annual rye) and Elymus elimoides (bottlebrush squirreltail) (3,000 seeds/m²) were used to remove nutrients from the soil. In another set of addition series matrices, nitrogen (33 kg/ha) or phosphorus (33 kg/ha) were added to the soil. Nutrient analysis of soil and vegetation indicated that annual rye and bottlebrush squirreltail reduced nutrient availability in soils. In another matrix, neither a background density nor nutrients were added. Data were fit into Watkinson's curvilinear model to determine the competitive relationship between bluebunch wheatgrass and spotted knapweed. This allowed comparison of the equivalence ratios (C) generated from each addition series. The C parameters are the per‐plant equivalent of bluebunch wheatgrass or spotted knapweed and can be interpreted as the ratio of intra‐to‐interspecific competition. The C parameters are also the equivalence ratio of the number of spotted knapweed it takes to have equivalent effect on bluebunch wheatgrass or the number of bluebunch wheatgrass having the equivalent effect on spotted knapweed. Without nutrient manipulation, spotted knapweed was more competitive than bluebunch wheatgrass. The C for bluebunch wheatgrass was 0.17, indicating that 0.17 knapweed plants were competitively equivalent to one wheatgrass. Annual rye changed the competitive balance in favor of bluebunch wheatgrass (C = 9.9). Addition of nitrogen, phosphorus, or the mid‐seral species did not change the competitive relationship between the two species. This preliminary study suggests that succession from spotted knapweed to late‐seral bluebunch wheatgrass community may be accelerated by altering resource availability.     

Cold hardiness in the scorpion, Paruroctonus aquilonalis.

Lower lethal temperatures represented by supercooling points were measured for the scorpion Paruroctonus aquilonalis (Stahnke) following exposure to laboratory regimes of photoperiod and temperature, as well as to ambient conditions. Analyses for the presence of glycerol and sorbitol in hemolymph, and of body water content were also performed.Laboratory results indicated that neither decreasing photoperiod nor decreasing temperature influenced supercooling point in a meaningful way. However, supercooling points did decrease significantly during the fall and winter among scorpions maintained outdoors, indicating development of cold hardiness. Neither glycerol nor sorbitol were detected, nor were the slight seasonal decreases found in water content considered sufficient to influence supercooling point.Ability to avoid winter freezing in P. aquilonalis appears primarily conferred by substantial seasonal depression of the supercooling point and is temporally associated with cessation of feeding in the fall. Supercooling responses were compared with those of another scorpion, Diplocentrus spitzeri, and differences tentatively were explained in terms of feeding activity.     

Effects of native species diversity and resource additions on invader impact

Theory and empirical work have demonstrated that diverse communities can inhibit invasion. Yet, it is unclear how diversity influences invader impact, how impact varies among exotics, and what the relative importance of diversity is versus extrinsic factors that themselves can influence invasion. To address these issues, we established plant assemblages that varied in native species and functional richness and crossed this gradient in diversity with resource (water) addition. Identical assemblages were either uninvaded or invaded with one of three exotic forbs: spotted knapweed (Centaurea maculosa), dalmatian toadflax (Linaria dalmatica), or sulfur cinquefoil (Potentilla recta). To determine impacts, we measured the effects of exotics on native biomass and, for spotted knapweed, on soil moisture and nitrogen levels. Assemblages with high species richness were less invaded and less impacted than less diverse assemblages. Impact scaled with exotic biomass; spotted knapweed had the largest impact on native biomass compared with the other exotics. Although invasion depressed native biomass, the net result was to increase total community yield. Water addition increased invasibility (for knapweed only) but had no effect on invader impact. Together, these results suggest that diversity inhibits invasion and reduces impact more than resource additions facilitate invasion or impact.     

Effect of summer drought relief on the impact of the root weevil Cyphocleonus achates on spotted knapweed

A recent decline in spotted knapweed, Centaurea stoebe L. subsp. micranthos (Asteraceae), has been observed in parts of western Montana. The release of the biological control agent Cyphocleonus achates (Fahraeus) is thought to contribute to the decline, but persistent drought since at least 1999 may be an additional factor. We conducted outdoor plot experiments to test the relative impacts of C. achates weevils and summer drought relief on spotted knapweed survival and growth. Groups of spotted knapweed transplants were assigned to one of four weekly water addition treatments (no added water, and 0.25, 0.5 or full recovery of plant water deficit, where "deficit" refers to potential evapotranspiration minus rainfall) in May to August 2004 and June to August 2005 and to either exposure to or protection from C. achates. In June of each subsequent year (2005 and 2006), plants were harvested and growth attributes that reflect plant vigor were measured. Drought indices showed that throughout the time of the study until January 2006, western Montana was in drought alert or severe drought. Summer drought relief had no effect on aboveground biomass and plant height of knapweed plants in subsequent years, but feeding by C. achates larvae reduced these two measures of plant vigor. Knapweed plants resuming growth after the drought ended in spring 2006 were significantly larger than those resuming growth under drought conditions in spring 2005. Spring drought may reduce knapweed growth, but C. achates reduced knapweed growth regardless of drought conditions.     

Distribution of Insect Attacks in Biological Control of Weeds: Infestation of Centaurea virgata Flowerheads by a Gall Fly

The success of biological control efforts to reduce weed density through release of insects may depend as much on the distribution of insect attacks among individual plants or plant parts as on the mean level of infestation. We used an index of dispersion to describe the distribution of Urophora quadrifasciata (Diptera: Tephritidae) galls among squarrose knapweed (Centaurea virgata) flowerheads at 18 west central Utah sites in the first 5 years following introduction of the biological control agent. Two thirds of the samples showed a significantly aggregated distribution of galls among flowerheads. Statistical analysis showed that site and year accounted for relatively small proportions of the variance in the index of dispersion. The degree of gall aggregation among flowerheads was positively correlated with the mean flowerhead quality (mean number of seeds per flowerhead; P = 0.013) and tended to be negatively correlated with the mean fly density per flowerhead at a site in a given year (P = 0.097). Our data suggest that higher quality flowerheads, and possibly higher quality plants, are preferentially attacked by U. quadrifasciata and therefore are more heavily subject to reduced reproductive potential through biological control. However, an aggregated distribution of fly attacks may undercut the potential of the fly to reduce seed production by the weed population as a whole. Understanding both the distribution of insect attacks among individual plants and the behavioral mechanisms producing such distribution patterns is important to the biological control of weeds.