RESEARCH Successful Biological Control of the Musk Thistle in Oklahoma Using the Musk Thistle Head Weevil and the Rosette Weevil M. Roduner, G. Cuperus, P. Mulder, J. Stritzke, and M. Payton ABSTRACT Rhinocyllus conicus Froelich, the musk thistle head weevil, was released in northeastern Oklahoma beginning in 1991 for biological control of Carduus nutans L., the musk thistle. By 2001, weevils had been released in 34 counties. Trichosirocalus horridus (Panzer), the rosette weevil, was released in six counties in 1998; and additional introductions in 2000 and 2001 brought the total number of county releases for this species to 29. Release areas were surveyed in 2000 and 2001 to determine the level of weevil establishment. Head weevils were recovered from 30 of 34 counties. Thistle populations were considered well infested if 30% of the heads had >4 larvae or pupae present. In 63% of the counties, ≥25% of the sites were well infested. Thistle densities had been reduced by 25–90% in 13 counties in Oklahoma where weevils had been released. Rosette weevils were recovered in three of the six original release counties and in one county where no releases were made until 2001. A combination of head and rosette weevils in the rosette weevil recovery areas provided a synergistic reaction with thistle density reductions occurring faster than in sites where only head weevils were released. Head weevils are established in Oklahoma and are effectively reducing thistle infestations. Rosette weevils also are established in several of the1998 release areas. C arduus nutans L., the musk thistle, is an exotic noxious weed that was accidentally introduced into the Unites States during the 1860s from Europe. Musk thistle has been present in Oklahoma since the 1940s. By 1960, it had spread through 29 counties in northeast and central Oklahoma (Stritzke et al. 1999); and by 2001, musk thistle was reported in 32 additional counties, thus 61 of 77 counties in the state were infested (Fig. 1). Thistle infestations are most frequently found in pastures, ditches, and abandoned areas. It displaces all other forms of herbaceous vegetation, reducing pasture yield. One musk thistle plant on 1.49 m2 (6,711 plants/ha) competing for space, light, and nutrients can reduce pasture yields by 23% (Trumble and Kok 1982). In addition, cattle refuse to feed on thistle plants because of the thorny stems and leaves (Rees 1991). The Oklahoma state legislature declared musk thistle a locally noxious weed (Northeastern Oklahoma) in 1994. In 2000, the legislature upgraded that status to encompass the entire state and provided specific rules on regulating density and control of the plants (Oklahoma Noxious Weed Law Rules 2000). Under these guidelines, a heavy density of musk thistle is 24 plants/ha, with a requirement that landowners prevent thistles from producing seeds. Landowners are considered out of compliance if there are >24 plants/ha or the plants are producing seed, or both. Once a landowner is determined to be out of compliance, fines are set at $1,000 per day. The law also gives counties the authority to spray herbicides on private property and apply those costs to a landowner’s property taxes. Because of unsatisfactory long-term control using chemical, cultural, and mechanical methods, the Oklahoma Cooperative Extension Service searched for new ways to control musk thistle. Kansas, Missouri, Texas, and Nebraska have successfully used biological 112 control (Fick and Peterson 1995, Wilson et al. 1996, Smith 2001, Jackman 2002a) to combat musk thistle infestations. Reductions in thistle stands in Missouri using Rhinocyllus conicus Froelich, the musk thistle head weevil, led to its introduction in Oklahoma during 1991. Subsequently, Trichosirocalus horridus (Panzer), the rosette weevil, was introduced into seven Oklahoma counties in 1998 to enhance control of musk thistle. In Virginia, Kok and Pienkowski (1984) used head weevils for long-term control of musk thistle, following thistle infestations from head weevil release to thistle stand collapse. Their 12-yr study in the same location showed a distinct pattern of thistle reduction. Head weevil larvae infest musk thistle seed heads and consume developing seeds and receptacle tissue. Each year less seed is produced, and fewer seeds are added to the soil seed bank. About 6 yr after weevil release, head weevil numbers were very large, and thistle stands collapsed in 10 yr (Kok and Surles 1975, Kok and Pienkowski 1984). Large numbers of weevils per head are not needed to significantly reduce seed production. One or two larvae per head causes some seed reduction (Surles and Kok 1978); four to five larvae per head leads to a 55% reduction in mature seed; and an increase to nine larvae per head can reduce mature seed by 98% (Rees 1991). Rosette weevils do not directly compete with head weevils. The rosette weevil larvae tunnel into plant roots feeding on the meristematic tissue causing sublethal damage. Feeding causes a black mass of frass and necrotic tissue in the rosette center; apical dominance is broken; and multiple short stems are produced with a reduction in heads and seed production. (Kok et al. 1986) (Fig. 2). Reduction in stands occurred in as few as 3–4 yr (Kok et al. 1986). Since 1991, when head weevil and rosette weevil were first released in Oklahoma (Stritzke et al. 1999), the only available inforAMERICAN ENTOMOLOGIST • Summer 2003 Fig. 1. Musk thistle locations in Oklahoma verified by survey. 1990 locations verified by Bill Stacey, Oklahoma Cooperative Extension Service; 2000 and 2001 locations verified by Oklahoma Cooperative Extension Service surveys and this study. mation pertained to where and when releases occurred. Although no data were taken initially on thistle stand size or density during release, there was abundant anecdotal evidence that head weevils were reducing thistle stands in the earliest release sites. The purpose of this study was to provide a detailed account of thistle infestations, and the population levels of head weevils and rosette weevils in 2000 and 2001. In this article, we compare survey data obtained during these 2 yr and anecdotal information with the thistle stand data of Kok and Pienkowski (1984) from Virginia. This information will help to establish a baseline for future musk thistle biological control in Oklahoma. Materials and Methods During the summers of 2000 and 2001, 36 counties were surveyed . Counties were chosen on the basis of head weevil releases that occurred from 1991 to 1999. Assessments followed several steps. We checked records from previous head weevil releases, and we contacted county Extension Educators for updated release lists. In areas where Extension Educators could not supply information, we contacted county road crews and Department of Agriculture employees for specific thistle information. We sent letters to landowners who had made releases to obtain permission to check thistle infestations on their land, and ≥50% responded. Musk Thistle Infestation. Musk thistle infestations were recorded by location using a gazetteer with the official Oklahoma road numbering system (DeLorme 1998). Levels of thistle infestation were compared with the levels in Oklahoma’s Noxious Weed Law Rules as follows: (1) Light infestation: <2 plants/acre (5/ha) (2) Medium: 2–9 plants/acre (5–24/ha) (3) Heavy: 10 or more plants/acre (25+/ha) (Oklahoma Noxious Weed Law Rules 2000). Any site >1 ha with musk thistle present was considered a large site, and we estimated the thistle density. These estimates were made with two transect lines used to count the number of plants per 305 m2. Head Weevil Detection and Density. At each location, 50 thistle heads were removed randomly throughout a thistle patch that constituted the sample area. Thistle patches ranged in size from 0.5 to 50 ha. Samples were bagged and dated, and location was noted. We also noted unusual situations that were encountered at each site including damage from other arthropods, wind, cattle, and any tillage or mowing operations. After visiting known release sites, we checked the surrounding area (within a 5-mile radius) in all directions by driving adjacent roads looking for additional thistle plants. Thistle heads were collected from these adjacent areas to assess the spread of weevil populations. Where information was not available Fig. 2. Rosette weevil damage, Rogers County, Okla., Bell Ranch. (top left) Rosette weevil damage in musk thistle rosette with necrotic tissue in the plant center. (bottom left) Musk thistle plants stunted by rosette weevils, ~550 mm; note multiple stems caused by larval damage, May 2002. (right) Normal musk thistle plant, ~1,450 mm. Photo of normal height plant located in Payne County, OK (May 2002) provided for comparison with rosette weevildamaged plants. There were no normal plants in the Bell Ranch pasture during 2001 or 2002 to use for comparison. AMERICAN ENTOMOLOGIST • Volume 49, Number 2 113 Fig. 3. Oklahoma counties with reductions in individual musk thistle stand densities ranging from 25 to 90%. Counties listed are those surveyed in 2000 and 2001. Numbered counties are referred to in the text. about weevil releases, or in counties bordering release areas, we traveled roads in a grid pattern to cover as much territory as possible. In 2000, 14 counties in northeastern Oklahoma were surveyed (Fig. 3). Digital photos were taken of known thistle densities and used to estimate densities in large pastures and fields. Photos and detailed maps were also used to verify thistle infestations. Large thistle infestations with head weevil infestations were rechecked in 2001. In 2001, 22 additional counties were surveyed, most in central and western Oklahoma. Both years’ thistle heads were brought back to the laboratory and frozen until they could be processed. Each head was measured in millimeters at the receptacle base, and the number of larvae, pupae, or pupal cases counted. In 2001, weevil-infested thistle heads were assessed based on the percentage of physical damage and the number of larvae, pupae, or pupal chambers recorded; 8,375 thistle heads were dissected in the laboratory, and 4,962 were infested with one or more larvae, pupae. All heads with four or more larvae and/or pupae per head were considered “well infested”. The percentage of well-infested heads in Fig. 4. Head weevil-infested thistle heads. Physical damage increases dramatically when numbers of pupae reach six and higher in all head diameters. (4,962 thistle heads analyzed by Sigma Plot [SPSS 1986– 1987]) 114 each sample of 50 heads was calculated. A site was designated as well infested if 30% of heads at a site contained four or more larvae and/ or pupae. The number of well-infested sites per county were divided by the total number of sites sampled to give the percentage of wellinfested sites in each county. Percentages of physical damage compared to weevils per thistle heads were analyzed using Sigma Plot (SPSS 1986–1987) to visualize the changes in damage levels (Fig. 4). This plot is a response surface plot of the means of the data reflecting the number of pupae and diameter of the thistle head. It is not based upon any model; therefore, no assessment of fit is given. These numbers were used to assess weevil establishment in each county. Any additional insects observed to cause damage to thistle seeds also were noted. The effects of these additional arthropods on thistle seed and/or plant production were assessed by counting the number of insects per plant or head and ultimately determining the damage to seed heads. Rosette Weevils. In 2001,we surveyed counties with rosette weevil releases using the same procedure as for head weevils. In early spring, before the flower stem elongated (bolted), plants were dug up and brought back to the laboratory to count the number of larvae in the meristematic tissue. Any rosette weevil adults captured outside known release areas were preserved as voucher specimens in the Oklahoma State University Research Collection to verify the spread of rosette weevils from previous release sites. Results During informal conversations, landowners and growers discussed their impressions of using head weevils for thistle control. Satisfaction with head weevils appeared to depend on the growers’ initial expectations. All growers who stated they were unable, physically or financially, to use herbicides were willing to allow weevils time to reduce thistle stands. Anecdotal reports were received that weevils began reducing thistle stand density in as little as 5 yr. Growers who were impatient or did not understand the thistle–weevil interactions, mowed thistles out of phase with plant and weevil growth, or applied herbicides after bolting and subsequently complained of slower reductions in their thistle stands. Musk Thistle Density. Musk thistle infestations in Oklahoma in 1990 covered 29 counties. In 2001, 61 counties were infested with this noxious weed. For this 2-yr study, initially, thistle density was counted as the number of mature (bolted) plants per 0.01 ha (≥305 m2). In 90% of the sites visited, thistle densities were greater than the numbers established as a heavy infestation (24 plants/ha) under the Oklahoma’s Noxious Weed Law. Some sites had thistle densities approaching 50,000 plants/ha (20,243 plants/acre). We made notations of areas with extremely heavy plant densities. Sites with heavy infestations of weak thistle plants (i.e., thin stemmed, few leaves, few seed heads, etc.) in 2000 either did not exist in 2001, or their densities were reduced dramatically. Reductions in thistle densities were observed, from as high as 20,000 plants/ha (2 plants/m2) in 2000 to as low as 10 plants/ha in 2001 (Fig. 5). Counties that experienced reductions in thistle density had 25–90% fewer plants. In three counties where rosette weevils were recovered, there was an additional reduction in thistle plant vitality. Thistle plants damaged by rosette weevils were short (average <0.5 m) compared with healthy plants (average 1.5m). Larvae consumed the main growth point, which broke apical dominance, causing plants to produce 3– 6 thin stems instead of one main stem. Damaged plants also had fewer heads (0–3) than healthy plants (5–35). Head Weevils. Large numbers of head weevil larvae were supported by thistle heads. When 15–25 larvae or pupae were present in a single head, it often appeared dry, shriveled, and brown (Fig. 6). AMERICAN ENTOMOLOGIST • Summer 2003 The percentage of physical damage increased rapidly when weevil levels exceeded 6 per head. In addition, 15 or more weevils caused 100% damage in heads 20 mm and larger (Fig. 4). Weevils were considered established at a release site (Fig 7A) if larvae, pupae, or empty pupal chambers were found inside thistle heads. Of the 36 counties surveyed (Fig. 7B), 2 had no present record of musk thistle, and 3 counties had thistle but no head weevils. All thistle heads checked in 7 counties had fewer than four individuals per head. Four counties had up to 24% of well-infested sites; in 6 counties 25–49% of all sites were well infested. Ten counties had 50–74% of well-infested sites, and 75–100% of sites in 4 counties were well infested. Thirteen counties had reductions of 25– 90% in thistle density between 2000 and 2001 (Fig. 5). Head weevils have been in these areas for a minimum of 8 yr. Thistle density reductions were especially dramatic in Okfuskee, Nowata, Craig, and areas of Cherokee, Delaware, and Adair Counties (Fig. 3). In 2000, nine sites in Okfuskee County (Fig. 3) were surveyed. Thistle densities were 2,500–7,500 plants/ha with >75% of the sites well infested with weevils. In 2001, these sites were sampled again, and 50–90% reductions in thistle density (3,750–250 plants/ha) were noted. The eastern border of Nowata County and western border of Craig County along US Hwy 60 (Fig. 3) contains several reclaimed mines. The reclamation land and surrounding areas have been heavily infested with musk thistle. Landowners describe a thistle infestation Fig. 5. Reduction of musk thistle at the Peabody Coal mine reclamation area, Craig County, OK, U. S. Hwy 60 and CR 210. (top) June, 2000. Plants weak with few blooms. (bottom) April, 2001, ~9 yr after head weevil infestation. Few plants or rosettes present. AMERICAN ENTOMOLOGIST • Volume 49, Number 2 Fig. 6. Comparison of uninfested and head weevil infested musk thistle heads. (left) uninfested with all seeds released (right) heavily infested, no seeds, head did not develop. >4,100 plants/ha from the 1980s into the early 1990s. Head weevils were released in this area in 1991 and, according to landowners, thistles have not been sprayed or mowed because of the excessive costs. Musk thistle density subsequently declined up to 90% in these areas. Thistle plants present in 2000 were thin and had only 3–4 heads per plant; rosettes were present in low numbers (Fig. 5A). In 2001, thistles were difficult to find at all of these sites (Fig. 5B). A large area encompassing the southern border of Delaware County and the northern borders of Cherokee and Adair Counties (Fig. 3) had thistle infestations of 24,700–37,000 plants/ha as reported by landowners. Head weevils were first released in Cherokee County in 1991 and Adair County in 1992, and then they were routinely released in subsequent years. In 2000, >50% of these sites were well infested by weevils. By 2001, reductions in the density of thistles from those recorded in 2000 varied from 30 to 75% by site with low numbers of rosettes present. Rosette Weevils. By 2001, 29 counties are known to have had rosette weevil releases (Fig. 8A.). In 2000 and 2001, rosette weevils were recovered from three of the seven counties where initial releases were made (Fig. 8B). During 1998, rosette weevils were released within 1 mile of Jay in Delaware County. During 2001, adult weevils were recovered in thistle patches from pastures along SR 127, at sites 0.5 and 1.5 miles west of US Hwy 59 (Fig. 8B). Approximately 60% of plants were short with multiple stems and reduced head numbers. In 1998, the first rosette weevil release in Rogers County was made at the Bell Ranch, north and east of Claremore (Fig. 8B). This site, ≥810 ha of reclaimed mine land, was infested at a thistle density of >50,000 plants/ha. In 1991, head weevils were released and provided good control. Rosette weevils were released in 1998 to augment the head weevil population. In 2000, ≥40% of the weevils collected for redistribution were rosette weevils. In 2001, ≥80% of the total weevils collected were rosette weevils. Adult weevil densities of up to 100 per plant were noted during collection. By 2001, the thistle-infested area was reduced to ≥16 ha with almost 100% of plants showing severe damage from rosette weevils (Fig. 2A). Adair County did not have an official rosette weevil release until 2000; however, adults were collected 2 miles south of Chewey, Okla. (Fig. 8B), several weeks before the rosette weevil release. Head weevil releases conducted in 1991–1993 were done with weevils collected in Missouri. An occasional container of head weevils had one or two adult rosette weevils, and this site likely received one of the contaminated containers (W. Stacey, personal communication, Oklahoma Cooperative Extension Service; Area Extension Entomology Specialist). If this is true, a significant rosette weevil population developed over 10 yr from relatively few weevils. The original release site has significant weevil damage, and the weevils are present (damaged 115 Fig. 7. (top) Head weevil release sites from 1991 through 2001. (bottom) Head weevil recovery sites in 2000 and 2001, showing the percentage of sites in a county with 30% or more of the heads per site infested with >4 larvae or pupae (L & P) per head. plants and adults recovered) in surrounding pastures within a 1mile radius. Other Insects. Three other insect orders were frequently observed in thistle heads. These species damaged thistle heads and affected seed production. In 2000, thistle heads from Cherokee and Delaware Counties had empty dipteran puparia. Several sites in each county had up to five pupae per head. The puparia were clear, blue or black, and empty, making identification impossible. Only occasional diptera puparia were present in 2001. During 2000, larvae of Homeosoma electellum (Hulst), the sunflower moth, (Lepidoptera: Pyralidae) were present in thistle heads in small numbers (1–2 larvae in 2–3% of heads). These numbers increased greatly in 2001, with 100% of secondary and tertiary heads infested in thistle patches in Payne County. The larvae fouled heads with their webbing and frass, making seed release difficult (Fig. 9A). Some of the smaller heads (8– 10 mm) were totally consumed. Euphoria sepulchralis (F.), flower scarab beetles, (Coleoptera: Scarabaeidae) were present in 2000–2001 in new blooms. Thistle heads were carefully cut apart to avoid disturbing the feeding beetles. Beetles were observed biting off the tops of developing seed. Previously damaged areas turned black and shriveled (Fig. 9B). Five to seven beetles were present in large thistle heads, buried head downward and actively feeding. Thistle heads with 5–7 beetles feeding had up to 75% of the seeds damaged. Discussion Head weevils have been successful in reducing musk thistle populations in several other states. Head weevils have been present in Virginia since the early 1970s with studies following the patterns of weevil increase and thistle reduction (Kok and Pienkowski 1984). Musk thistle in western Missouri has been reduced to the point that weevil collectors from Oklahoma were unable to find enough weevil numbers for redistribution (W. Stacey, personal communication). Missouri, Nebraska, and Kansas have active biological control programs collecting head weevils and releasing them in new areas. All of 116 these states have areas with successful control of the musk thistle (Fick and Peterson 1995, Wilson et al. 1996, Smith 2001). In Oklahoma, head weevils are associated with reduced musk thistle densities where they have been established for longer than 6 yr. Since the release program began, 13 counties have experienced reductions of thistle stands (Fig. 3). Weevils were recovered from 30 of 34 release counties and 25 counties have sites that are considered well infested (Fig. 7B). Head weevils have spread from their original release sites infesting thistles in surrounding areas, generally following the prevailing winds. This pattern is similar to that observed in Virginia (Kok and Surles 1975). Counties with only one or two known release sites had weevils scattered throughout the county. In Payne County, for example, head weevils were released on two sites in the early 1970s and are now established throughout the county. Thistle infestations were conspicuously absent in areas with dense grasses, healthy pastures, or heavy shade. The majority of infestations occurred in ditches, pastures with weak stands of grass, disturbed or abandoned areas, and hard-red winter wheat fields. Land with gullies or water courses that either facilitated seed movement or trapped it had extremely heavy thistle infestations. Reductions in thistle size and vitality, with head weevil infestation and dense pastures were observed in previous studies done in Virginia. Researchers followed thistle seed consumption by head weevils. The highest quality seed was consumed first, leaving lower quality seed to enter the soil seed bank. As the vitality of thistles decreased, pastures were able to recuperate, competing well with thistles for space and nutrients. This competitive effect appeared to be synergistic resulting in a “crash” of the thistle population (Kok and Pienkowski 1984, Kok et al. 1986). Locations in Oklahoma where weevils have been present for 6– 10 yr resemble this scenario. Thistle plants in these areas were thinner and had fewer heads than thistle plants in areas where weevils were not present. Although no data on thistle density were taken at the time of weevil releases, by comparing what landowners reported as thistle density at the time of these releases with the reductions observed Fig. 8. (top) Rosette weevil release sites from 1998 to 2001. (bottom) Rosette weevil recovery sites from 2000 to 2002. Numbered counties are referred to in text. AMERICAN ENTOMOLOGIST • Summer 2003 Fig. 9. Damage from other insects: (left) Webbing and frass caused by the sunflower moth fouling a thistle head. (right) Individual musk thistle seed damaged by the flower scarab beetle. from 2000 to 2001, the time frame and pattern of thistle reduction follow the results from Virginia reported by Kok and Pienkowski (1984). Head weevil populations increased, and the thistle populations collapsed about 10 yr after the weevils were introduced. Low numbers of rosette weevils follows the pattern of seed reduction outlined by Surles and Kok (1978) and Rees (1991) because fewer seeds were available to germinate in the soil seed bank. Rosette Weevils. Adding rosette weevils to locations already infested with head weevils provides additional control (Cartwright and Kok 1985, Kok et al. 1986). Recovery areas for rosette weevils in Adair, Delaware, and Rogers Counties all had head weevils present first. When rosette weevil populations began to increase, thistle populations dropped faster than with head weevils alone, verifying the results of Cartwright and Kok (1985) and Kok et al. (1986). Because the weevils occupy different feeding niches, there appeared to be no direct competition. Rosette weevils were first collected in northern Kansas in 1998 for release in Oklahoma. In 2000, weevils were collected from the state of Kansas and Rogers County, Okla., and were released in 22 counties in Oklahoma. In 2001, weevils were collected in Rogers County and distributed to additional locations in counties that had earlier releases and at least three additional counties. Rosette weevils increased so rapidly that one site supplied adequate numbers for new release locations without reducing effectiveness in that area. Personnel from several counties in western Oklahoma shared weevils with other counties. Although no data on thistle stand density or size were taken during these initial releases, in the three counties where they have become established, rosette weevils in combination with head weevil populations appear to have caused musk thistle decline faster than in pastures with head weevils alone. The Bell Ranch in Rogers County provides overwhelming evidence of this phenomenon. The landowner reported thistle densities of ≥50,000 plants/ha in 1991 when the first head weevils were released. In 1998, he estimated that thistle plants had been reduced by >50% (<20,000 plants/ha). This was the same year that rosette weevils were first released into the site. In 2002, 4 yr after releases of rosette weevil, thistle density stood at <15 plants/ha, and those plants remaining did not develop viable seed heads. The county Extension Educator reported that all rosettes examined during the spring of 2002 contained several hundred rosette weevil larvae. Surviving plants averaged <0.3 m tall and had multiple stems and few heads (Fig. 2B). Rosette weevils have spread into neighboring areas and were collected in large numbers from these sites for redistribution. In 2003, the Bell Ranch along with most of Rogers County, Nowata County, and Okfuskee County are no longer viable locations for collection of weevils for the Oklahoma AMERICAN ENTOMOLOGIST • Volume 49, Number 2 Musk Thistle Roundup because of the low population of thistles throughout the area (Pratt 2003). Other Insects. Three other insects were found in thistle heads that had some effect on thistle seed production. Dipteran larvae destroyed some receptacle tissue and associated seeds. Sunflower moth larvae, although pests of commercial sunflower fields, also provided thistle seed reduction. Infestations of sunflower moths have been noted in other areas of the country, and their effectiveness was related to infestation levels (Goyer 1978). Population densities of sunflower moths vary from year to year, and they can be considered beneficial, if they do not affect nearby cash crops (McCarty 1982). Flower scarab beetles also were present in thistle heads in numbers large enough to be noticeable. In addition to reducing the amount available for pollination and consuming nectar and pollen (Hayes 1929), all three additional insect species fed on newly developing seeds, causing severe damage and preventing seed maturation. Although anything that destroys developing seed could be considered beneficial, damage on the plant did not appear to be consistent enough to provide reliable control. Additional studies will be needed to determine if any of these insects are viable and reliable candidates for biological control releases. Need for Vigilance. Actual musk thistle densities are much higher than those described in the Oklahoma Noxious Weed Law Rules. According to these criteria, the legal definition of high thistle density is 10 plants/0.4 ha (1 acre) or 1 plant/404.6 m2 (4,356 ft2). Thistle density in several locations was as high as 50,000 plants/ha (20,243 plants/acre) or 1 plant/0.2 m2 (2.2 ft2). Grower education about musk thistle growth patterns and its ability to reproduce explosively is of paramount importance. It is easy to discount densities as low as those in the Noxious Weed Law, but thistles must be controlled when populations are low to prevent heavier infestations that require increased time for beneficial organisms to assume a regulatory role. Head weevil population increases are rapid (Kok and Surles 1975, Rees 1991), so sites that were considered well infested (30% of heads with >4 larvae or pupae per head) would be heavily infested with weevils within 1 or 2 yr. Rosette weevils have experienced a greater rate of increase than head weevils, especially at the Rogers County site. Counties with head weevils and rosette weevils can anticipate rapid reductions of thistle infestations. This snapshot of musk thistle biological control taken during 2000–2001 shows a pattern similar to the one observed by Kok and Pienkowski (1984) during their long-term studies of biological control in Virginia. Although previous data for Oklahoma is unavailable, anecdotal evidence throughout the state is consistent. Musk thistle densities were much higher before the introduction of head and rosette weevils, and in the 10 yr since the release program began, thistle densities are now lower. Landowners who have given weevils a chance to work are not only pleased with the results but are telling others about their success. Requests for weevil releases are increasing each year as landowners see the results in neighboring areas. The head and rosette weevil release program has been a resounding success in northeastern and central Oklahoma. With 1 musk thistle plant on 1.49 m2 (6,711 plants/ha) reducing pasture yields by 23% (Trumble and Kok 1982), and many thistle stands three to four times more dense, significant thistle reduction will decrease costs associated with land management. Economic Benefits. Based on a survey of Oklahoma growers, the average amount of improved pasture for each producer ranges from 16 to 64 ha (New 1997). The average cost of controlling musk thistles for a year using 2,4-D herbicide is estimated at a minimum of $1.85/ha for 0.39 liters of material per ha (not including application costs) (Medlin et al. 2003). There are about 7.1 million acres of 117 improved pastures in Oklahoma. Therefore, the statewide cost of controlling musk thistle with herbicides for 10 yr, if all improved pastures were infested, would exceed $131 million. Substantial savings are possible when musk thistle weevils are integrated into a musk thistle management system. Spraying of pastures would be phased out after 3 yr, and no annual border spraying would be required. Costs associated with an integrated approach (still relying on herbicide application for the initial 3 yr) using weevils would be $5.55/ha for spraying over the 3 yr and $3.45/ha associated with trips to collect weevils. This represents a savings of at least $67 million over the first 10 yr, while at the same time reducing the amount of herbicide broadcast into the environment. Up to this point, the estimated costs ($3.45/ha) associated with weevil collections in Oklahoma have been provided by Extension Educators and the Integrated Pest Management Program at Oklahoma State University at no cost to participating producers. Not without Controversy. There is currently, some controversy in adjoining states about the use of head weevils for thistle control. This controversy relates to several species of Cirsium thistles being attacked by the head weevil. These thistles include the Platte thistle (C. canescens), Pitcher’s thistle (C. pitcheri), and wavyleaf thistle (C. undulatum) (Louda 1998, 1999). In addition, head weevil infestations have been recorded from the milk thistle (Silybum marianum; Jackman 2002b). Presently, from this list of thistles, only the wavyleaf thistle is found in Oklahoma (Stritzke and Tyrl 2000). Wavyleaf thistle heads were checked during this study when they appeared to be at an appropriate stage to support head weevils. Little to no survival of weevils was recorded on this species of thistle. It is unknown whether the hotter, drier climate in Oklahoma affects head weevil survival in nonhost species. Thistle stands in Oklahoma can be so extensive (up to 50,000 plants/ha and 100+ ha in size) that all other plants are excluded, and 118 AMERICAN ENTOMOLOGIST • Summer 2003 the land becomes unusable. With infestations of this magnitude and only one other possible host present, the use of head weevils for biological control has not been a serious issue. Members of the release teams and the Oklahoma Cooperative Extension Service are aware of these concerns; however, presently the benefits of using head weevils for musk thistle control far outweigh the concerns. They have also suggested that this question be revisited if other Oklahoma thistle species become more common and are shown to be in danger. In summary, the head weevil, has become established in Oklahoma, especially where it has been in place for at least 4 yr. During that time, musk thistle infestations have been reduced in density and plant vigor. Furthermore, the addition of the rosette weevil is increasing the speed of that reduction in thistle infestation. Efforts must not stop at this point. Thistle populations in the western half of Oklahoma have had weevils for only 1 or 2 yr and need time to establish. Continued releases of head weevil and rosette weevils and grower education to manage them will reduce thistles in the remainder of the state, allowing growers profitable use of their land. Acknowledgments The authors thank all the landowners and extension personnel who provided information about thistle infestations. We extend our sincere gratitude to laboratory technicians Tererai Nyamanzi, Audrey Sheridan, and Penny Potter for processing thousands of thistle heads. We are grateful to USDA–APHIS for all the survey and weevil release work they have supported in previous years that brought us to this point. We also thank our preliminary reviewers Patricia Bolin and Thomas Phillips of the Department of Entomology and Plant Pathology at Oklahoma State University for their helpful comments when reviewing this manuscript. This work was approved for pub- lication by the director of the Oklahoma Agricultural Experiment Station, and supported in part under project OKLO2173. This manuscript is part of a thesis submitted by M. Roduner to Oklahoma State University. References Cited Cartwright, B., and L. Kok. 1985. Growth responses of musk and plumeless thistles (Carduus nutans and C. acanthoides) to damage by Trichosirocalus horridus (Coleoptera: Curculionidae). Weed Sci. 33: 57–62. DeLorme. 1998. Oklahoma Atlas and Gazetteer. DeLorme Publications, Yarmouth, ME. Fick, W. D., and D. Peterson. 1995. Musk thistle identification and control. L-231. Cooperative Extension Service, Kansas State University. Goyer, R. 1978. Occurrence of the sunflower moth Homeosoma electellum (Lepidoptea: Pyralidae) on musk thistle in Louisiana. La. Acad. Sci. 41: 16–17. Hayes, W. P. 1929. Morphology, taxonomy, and biology of larval Scarabaeoidaea. Ill. Biol. Monogr. 12(2):10. Jackman, J. A., P. Boldt, J. W. Stewart and T. W. Fuchs. 1992. Biological control of weeds in Texas, musk thistle. http://insects.tamu.edu/extension/bullentins/1-5067.html. Kok, L. T., and R. L. Pienkowski. 1984. Biological control of musk thistle by Rhinocyllus conicus (Coleoptera: Curculionidae) in Virginia from 1969 to 1980, pp. 433–438. In E. S. Delfosse [Ed.]. Proceedings, 6th International Symposium on Biological Control of Weeds. Minister of Public Works and Government Services of Canada, Vancouver, British Columbia Agriculture Canada, Ottawa. Kok, L. T., and W. W. Surles. 1975. Successful biocontrol of musk thistle by an introduced weevil, Rhinocyllus conicus. Environ. Entomol. 4: 1025–1027. Kok, L. T., T. McAvoy, and W. Mays. 1986. Impact of tall fescue grass and Carduus thistle weevils on the growth and development of musk BIOSERV Artwork provided with proofs AMERICAN ENTOMOLOGIST • Volume 49, Number 2 119 thistle (Carduus nutans). Weed Sci. 34: 966–971. Louda, S. 1998. Population growth of Rhinocyllus conicus (Coleoptera: Curculionidae) on two species of native thistles in prairie. Environ. Entomol. 27: 834–841. Louda, S. M. 2000. Negative ecological effects of the musk thistle biological control agent; Rhinocyllus conicus, pp.215-243. IN Follet, P.A. and J.J. Duan, (eds.) Nontarget effects of Biological control. Kluwer Academic Publishers, Boston, MA, USA. McCarty, M. 1982. Musk thistle (Carduus thoermeri) seed production. Weed Sci. 30: 441–445. Medlin, C., P. Bolin, M. Roduner, L. Cargill, and P. Mulder. 2003. Integrated management of invasive thistles in Oklahoma. Oklahoma Cooperative Extension Service Fact Sheet F-7318, Oklahoma State University, Stillwater OK. New, M. G. 1997. Survey of weed management practices in pastures and rangelands in Oklahoma and selectivity of various herbicide treatments on cultivars of forage bermudagrass (Cynodon dactylon). M.S. Thesis, Oklahoma State University, Stillwater. Oklahoma Noxious Weed Law Rules. 2000. Title 35. Oklahoma State Dept. of Agriculture Chapter. 30. Plant Industry Subchapter. 34. Noxious Weeds § 3-220. Noxious Weeds—Eradication. Pratt, P.W. 2003. Musk thistle weevil roundup. Oklahoma Cooperative Extension Service, Timely Topics Newsletter 5:4, Oklahoma State University, Stillwater OK. Rees, N. 1991. Biological control of thistles, pp. 264–273. In L. F. James [Ed.]. Noxious range weeds. Westview Press, Boulder CO. Smith, T. 2001. Vegetation management guide, musk thistle. http:// www.conservation.state.mo.us/nathis/exotic/vegman/eighteen.htm SPSS. 1986–1987. Sigma Plot for Windows, Version 4.0, 233 S. Wacker Drive, 11th Floor. Chicago, Ill., 60606-6307. Stritzke, J., and R. Tyrl. 2000. Thistles in Oklahoma and their identification. Oklahoma Cooperative Extension Service. Fact sheet F-2776, Oklahoma State University, Stillwater, OK. Stritzke, J., B. Stacey, and G. Cuperus. 1999. Integrated control of musk thistle in Oklahoma. Oklahoma Cooperative Extension Service. Fact sheet F-7318, Oklahoma State University, Stillwater, OK. Surles, W. W., and L. T. Kok. 1978. Carduus thistle seed destruction by Rhinocyllus conicus. Weed Sci. 26: 264–269. Trumble, J. T., and L. T. Kok. 1982. Integrated pest management techniques in thistle suppression in pastures in North America. Weed Res. 22: 345–359. Wilson, R., F. Roeth, and A. Martin. 1992. Musk Thistle. Cooperative Extension, Institute of Agriculture and Natural Resources. NebGuide G1109-Weeds A-30, Field and Pasture. University of Nebraska-Lincoln. http://www.ianr.unl.edu/pubs/weeds/g1109.htm. Mary A. Roduner has completed her degree requirements for a Mas ters of Science in Entomology at Oklahoma State University and iscurrently located in Saba, Netherlands Antilles in the Caribbean, where her husband is attending medical school. Gerrit W. Cuperus is currently on disability leave from Oklahoma State University where he serves as Professor and IPM Coordinator for the Division of Agricultural Sciences and Natural Resources. His research interests are related to IPM in stored products. Phillip G. Mulder, Jr. is Extension Entomologist and Associate Professor in the Department of Entomology and Plant Pathology at Oklahoma State University. His extension/research interests focus on IPM in alfalfa, fruit trees, pecans, peanut, soybean, grapes and 4-H youth programs. In addition, he serves as Extension Coordinator for the entomology portion of the department and President-Elect of the Southwestern Branch of the ESA. Jimmy F. Stritzke is Professor Emeritus within the Department of Plant and Soil Sciences at Oklahoma State University. His research/extension interests are in the area of management of native and improved pastures. He retired from the Oklahoma Cooperative Extension Service in June, 2001. Mark E. Payton, is a Professor within the Department of Statistics at Oklahoma State University. He currently teaches and serves as statistical consultant for the Division of Agricultural Research and Natural Resources. 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