Rhinocyllus conicus (thistle-head weevil)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Air Temperature
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Rhinocyllus conicus Froelich 1792
Preferred Common Name
- thistle-head weevil
Other Scientific Names
- Curculio conicus
International Common Names
- English: thistle head weevil
Local Common Names
- Australia: receptacle weevil; seed-head weevil
- New Zealand: nodding thistle receptacle weevil
- RHILCO (Rhinocyllus conicus)
Summary of InvasivenessTop of page
Native to Europe and western Asia, R. conicus has been deliberately introduced to Canada in 1968 (Harris and Zwölfer, 1971; Harris, 1984); South America in 1980 (Feldman, 1997); Australia in 1989 (Woodburn and Cullen, 1993; 1995); and New Zealand in 1973 (Jessep, 1975; 1981) as a biological control agent for thistles in the genera Carduus,Cirsium and Silybum. Introduced populations were collected from different hosts and so included populations from both temperate and Mediterranean climates.
North America has a number of species of native thistle in the genus Cirsium that are susceptible to damage from R. conicus. As the impacts of R. conicus on some of them has been quite significant (Louda et al., 1990; 1995; Louda and Potvin, 1995; Louda 1998), and as some of these native thistles were already threatened, R. conicus is now considered an invasive species within parts of North America (Louda et al., 1997). At the time of its release, the potential impact of R. conicus on native thistles was considered acceptable.
In regions other than North America, no vulnerable native or economically important plants occur within the host range of R. conicus. It is not a regulated pest in any country of introduction.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Curculionidae
- Genus: Rhinocyllus
- Species: Rhinocyllus conicus
Notes on Taxonomy and NomenclatureTop of page
Rhinocyllus conicus is a weevil, native to Europe and western Asia, that has been introduced into several countries as a biological control agent for some exotic invasive thistle species (Boldt and Kok, 1982). Rhinocyllus is a small genus of five species in the tribe Rhinocyllini. Rhinocyllini contains only one other genus, Bangasternus, which contains seven species (Zwölfer and Harris, 1984). All weevils from both genera are believed to feed in the capitula of Asteraceae species. R. conicus has eight synonyms (Hoffmann, 1956; Alonso-Zarazaga and Talamelli, 2011).
Allozyme and morphometric analysis of R. conicus, from the Atlantic coast to Israel, found two distinct groups (Klein and Seitz, 1994). The two groups, temperate and Mediterranean, correspond to differences in climate, and differ in their phenology of oviposition and physical appearance: the Mediterranean group starts to lay about a month earlier than the temperate group and generally has a narrower body shape, although there is individual variation and overlap between the two groups. On this basis, and given the genetic divergence between groups, Klein and Seitz (1994) argued that R. conicus has two subspecies. They also argued that the Mediterranean group matches the morphological descriptions and climatic distributions of R. oblongusCapiomont (1873) and therefore that the status of this species needs further consideration.
DescriptionTop of page
Eggs are laid under frass caps on the external surface of the bracts of the pre-flowering capitulum. The four instars of larval development and pupation occur inside the capitulum. The adult is a typical ‘snout-beetle’, 6 (3-7) mm long. It is dark in colour with many clusters of vertical short, brown hairs that give it a ginger-speckled appearance. The hind wings are well developed as the insect is a strong daylight flier (Jessep, 1981).
DistributionTop of page
The known distribution of R. conicus does not perfectly match the distribution of its main host plants in its native range, which suggests some apparent gaps in its native range may simply be because the host has been recorded but the pest has not. R. conicus is likely to have a largely continuous distribution around the Mediterranean corresponding to the distributions of its main host plants (see Hosts/Species Affected).
Its true range in North America is likely to be greater than what has been recorded. Zwölfer and Harris (1984) stated that R. conicus ‘has been established in most parts of North America with a Carduus nutans problem’; C. nutans is known from nearly all of North America except 5 US states and 4 Canadian Provinces (USDA, 2013).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Greece||Present, Localized||Native||Mainland & Crete|
|Italy||Present, Widespread||Native||Including Sicily & Sardinia|
|Russia||Present||Present based on regional distribution.|
|United States||Present||Present based on regional distribution.|
|-California||Present, Widespread||Introduced||1971||Goeden 1995 for release date|
|-Colorado||Present, Widespread||Invasive||Based on impact on native thistles (DePrenger-Levin et al. 2010)|
|-Nebraska||Present, Widespread||Introduced||Invasive||Based on impact on native thistles (Louda et al. 1990)|
|-New Jersey||Present, Widespread||Introduced|
|-New York||Present, Widespread||Introduced|
|-North Carolina||Present, Widespread||Introduced|
|-North Dakota||Present, Widespread||Introduced||1975|
|-Oregon||Present, Widespread||Introduced||1978||Goeden 1995 for release date|
|-South Dakota||Present, Widespread||Introduced|
|-Tennessee||Present, Widespread||Introduced||Invasive||Based on impact on native thistle in controlled studies, not natural populations (Wiggins et al. 2010)|
|-Washington||Present, Widespread||Introduced||First reported: 1980s|
|-West Virginia||Present, Widespread||Introduced|
|-Wisconsin||Present||Introduced||1975||Based on impact on native thistle (Sauer and Bradley 2008)|
|Australia||Present||Present based on regional distribution.|
|-New South Wales||Present, Widespread||Introduced|
|New Zealand||Present, Widespread||Introduced|
History of Introduction and SpreadTop of page
R. conicus was first considered as a biological control agent for the exotic nodding thistle Carduus nutans by Canada. In 1968 the weevil was introduced to Saskatchewan and Ontario from Alsace, France, and the Rhine valley, Germany. The weevil proved an effective agent against C. nutans and also attacked another exotic plumeless thistle, C. acanthoides. From Saskatchewan and Ontario R. conicus was moved and released into the USA, in Virginia and Montana, in 1969. Again it proved effective against the target and was spread widely throughout the country.
Populations of the weevil from Italy, which may be a separate ecotype (Zwölfer and Harris, 1984) or subspecies (Klein and Seitz, 1994), were released in California on both variegated or milk thistle Silybum marianum and slender or Italian thistle Carduus pycnocephalus in 1971 and 1973 respectively, but proved ineffective (Goeden and Ricker, 1977; 1978).
New Zealand imported and released weevils from Canada in 1973, which proved effective against C. nutans in New Zealand.
Argentina released weevils from the USA in 1980, largely against C. acanthoides, a plant which has caused considerable damage in Argentina (Feldman, 1997).
In 1989 Australia released three populations of R. conicus, from New Zealand, southern France and Italy, into the state of New South Wales to counter C. nutans there (Woodburn and Cullen, 1993; 1995; Cullen and Sheppard, 2012), but only the New Zealand and French populations spread widely. Overall, R. conicus was less effective in Australia due to asynchrony with its host. Populations of the weevil from south-west France were released against spear thistle Cirsium vulgare in the state of Victoria in Australia in 1989, but establishment has been poor since (Sagliocco et al., 2012).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Australia||France||1989||Biological control (pathway cause)||Yes||No||Woodburn and Cullen (1995)|
|Australia||New Zealand||1989||Biological control (pathway cause)||Yes||No||Woodburn and Cullen (1995)|
|Canada||France||1968||Biological control (pathway cause)||Yes||No||Harris (1981)||Effective agent|
|Canada||Germany||1968||Biological control (pathway cause)||Yes||No||Harris (1981)||Effective agent|
|New Zealand||Canada||1973||Biological control (pathway cause)||Yes||No||Jessep (1981)||Effective agent|
|USA||Canada||1969||Biological control (pathway cause)||Yes||No||Andres and Rees (1995)||Effective agent|
|USA||Italy||1969||Biological control (pathway cause)||Yes||No||Andres and Rees (1995)||Effective agent|
Risk of IntroductionTop of page
The risks of the introduction and spread of R. conicus to other regions is limited, as it is specific to a few genera of host plants and has only left its native range through deliberate release as a biological control agent. Further deliberate introductions seem unlikely as there are few other countries where the thistle host plants are serious weeds.
The negative impacts of this insect are restricted to North America as this is the only continent with native (or commercial) plants that are susceptible to attack. R. conicus is probably already in most parts of the USA where its target hosts grow, so risks of further spread within the USA are only to regions with susceptible native thistles but without any of the relevant exotic thistles. To reduce risk of spread within the USA, R. conicus is now banned from inter-state movement.
HabitatTop of page
R. conicus is restricted to the habitats and regions where its host plants are found (see Hosts/Species Affected). The host plants are found in pastures and disturbed sites, like roadsides, in temperate and Mediterranean climates in Europe, North Africa, Western Asia, North America, southern South America, south-eastern Australia and New Zealand.
Habitat ListTop of page
|Terrestrial||Managed||Managed grasslands (grazing systems)||Principal habitat||Natural|
|Terrestrial||Managed||Industrial / intensive livestock production systems||Secondary/tolerated habitat||Natural|
|Terrestrial||Managed||Disturbed areas||Secondary/tolerated habitat||Natural|
|Terrestrial||Managed||Rail / roadsides||Secondary/tolerated habitat||Natural|
|Terrestrial||Managed||Urban / peri-urban areas||Secondary/tolerated habitat||Natural|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Secondary/tolerated habitat||Natural|
|Terrestrial||Natural / Semi-natural||Riverbanks||Secondary/tolerated habitat||Natural|
Hosts/Species AffectedTop of page
North America has about 90 species of native thistle in the genus Cirsium (USDA, 2013). Susceptibility of these species to R. conicus is related to a) their proximity to exotic host thistle populations on which this weevil is found, and b) the degree to which flowering phenology is synchronous with the reproductive cycle of R. conicus (Russell et al., 2007). A number of rare and threatened native Cirsium species in North America have been documented: these include C. canescens Nutt. (Louda et al., 1990; Arnett and Louda, 2001), C. undulatum Nutt. (Maw, 1982), C. ownbeyi S.L. Welsh (DePrenger-Levin et al., 2010) and C. hillii (Canby) Fernald (Sauer and Bradley, 2008). Turner et al. (1987) listed another twelve species of native Cirsium thistles in California from which R. conicus has been reared. Maw (1982) also found eggs on the native C. flodmanii (Rydb.) in Canada.
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
R. conicus lays frass-covered eggs on the exterior involucral bracts of immature inflorescences of its host. The larvae hatch and bore into the receptacle of the capitulum, destroying the reproductive surface from which achenes develop. A single larva destroys on average 26 seeds (Sheppard et al., 1994).
List of Symptoms/SignsTop of page
|Fruit / internal feeding|
|Inflorescence / dieback|
|Inflorescence / dwarfing; stunting|
|Inflorescence / internal feeding|
|Seeds / discolorations|
|Seeds / empty grains|
|Seeds / fused together|
|Seeds / shrivelled|
|Whole plant / internal feeding|
Biology and EcologyTop of page
The various groups and ecotypes of R. conicus have been described but not been fully understood genetically. Some may be subspecies. Populations from different regions and hosts have distinct host preference profiles and capacities to differentially exploit different thistle hosts (C. nutans, C. pycnocephalus, C. vulgare (Savi) Tenore and Silybum marianum Gaert.) (Unruh and Goeden, 1987). Populations from Onopordumacanthium have also been described (Zwölfer and Harris 1984). More work is needed to better understand the genetic patterns underlying these differences and to see if the different ecotypes show any degree of reproductive incompatibility.
R. conicus is a univoltine to partly bivoltine weevil. Adult females lay about 200 (54-360) eggs which take about a week to hatch. There are four larval instars which feed and pupate internally in the capitulum, in a hard black ovoid cell that resembles a gall. Pupation takes up to two weeks but the immature adults stay in the cells several weeks. Development from egg to adult takes about seven weeks, but may be shorter in Mediterranean climates. Newly emerged adults seek aestivation sites in summer in Mediterranean regions, whereas in temperate regions a second partial generation takes place. In late summer and in autumn the adults seek overwintering sites away from their hosts. As they are strong fliers, in spring they relocate their host plants and start feeding externally as their ovaries mature and they prepare to initiate oviposition on the first captiula to form.
Physiology and Phenology
This weevil only has a partial second generation in more temperate climates (Gassmann and Kok, 2002); however, in Tennessee and Georgia, R. conicus was observed to complete one generation and then not feed on thistles until the following spring (Wiggins, 2013, personal communication). Zwölfer and Harris (1984) considered that day-length determines second generations, so that new adults experiencing increasing daylength will start a second generation on flower heads of their host (mainly Carduus nutans) which developed later in the season. The first generation is very much skewed towards host captiula produced in spring (Cullen and Sheppard, 2012), which are also the largest; this means that later, smaller capitula receive proportionally fewer eggs. The capacity for a second generation is assisted by a longer flowering period of the host in temperate wetter summers.
In the Mediterranean region, where R. conicus also uses the hosts C. pycnocephalus, C. tenuiflorus and S. marianum, the hot dry summer means the hosts have a relatively early, but shorter flowering period, meaning R. conicus starts ovipositing earlier than in temperate regions, but only has time for one generation before their hosts stop flowering and adults need to seek aestivation sites. These differences relate to the different ecotypes by both region and host, and allow this species to persist under a broader range of hosts and climates.
In the exotic range these differences in phenology by ecotype are maintained with little evidence of quick synchronization to new conditions (Cullen and Sheppard, 2012), suggesting the ecotypes are phonologically distinct and not a phenotypically plastic strategy of survival. Effectiveness as a biocontrol agent and the impact of R. conicus on native thistles in the exotic range are determined by the degree of synchrony between the weevil oviposition and flowering of available potential hosts (Goeden and Ricker, 1985). Therefore the native thistles most impacted on by R. conicus tend to be those that flower early in the growing season. As a biological control agent it has proved more effective in temperate regions and this may be in part because the populations of the weevil introduced into these regions have a partial second generation.
Adults live up to 12 months.
Mediterranean-adapted populations undergo aestivation through the dry summer months. All populations hibernate through the cold months and migration tends to occur in spring, when the adults seek host populations having emerged from their hibernation sites. Once the adults have located host populations they become much more sedentary.
Population Size and Density
Population size is determined by the abundance of their principal host plants, but can range up to tens of individuals per square meter in spring when there are dense populations of host plants.
Nutrition is restricted to known host plants in the genera Carduus, Cirsium, Silybum and Onopordum.
This weevil is one of a diverse community of insects found in the capitula of its host plants in its native range (Sheppard et al., 1991; 1994). Other insects in this community tend to be active later in the flowering periods, but competition in the captula is quite intense. In the exotic range, the insect community is largely limited to other species that have been introduced as biological control agents in the exotic hosts (e.g. Woodburn, 1996); however, amongst the native thistles R. conicus competes in the developing capitula with native insects, particularly tephritid seed flies, and may locally threaten the population viability of such flies (Tatyana and Louda, 2011).
ClimateTop of page
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Tolerated||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
|Df - Continental climate, wet all year||Preferred||Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)|
|Ds - Continental climate with dry summer||Preferred||Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)|
|Dw - Continental climate with dry winter||Tolerated||Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)|
Air TemperatureTop of page
|Parameter||Lower limit||Upper limit|
|Mean maximum temperature of hottest month (ºC)||25||40|
|Mean minimum temperature of coldest month (ºC)||-20||10|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Bracon urinator||Parasite||Arthropods|Larvae||not specific||Native range||N|
|Exeristes roborator||Parasite||Arthropods|Larvae||not specific||Native range||N|
|Macroneura vesicularis||Parasite||California||Carduus nutans|
|Microdontomerus anthonomi||Parasite||USA; California||Carduus nutans|
|Neocatolaccus tylodermae||Parasite||USA; California||Carduus nutans|
|Pterandrophysalis levantina||Parasite||Eggs||to species||Native range||N|
Notes on Natural EnemiesTop of page
R. conicus is attacked by many different species of parasitoid in its native range (Zwölfer and Harris, 1984). The impact of these parasitoids on native R. conicus populations can be very significant, with particularly high levels of egg parasitism. Indeed, it appears that R. conicus lays more eggs on early capitula than the individual captulum can support, as in the native range parasitoids often kill a proportion of eggs; however, this over-laying of eggs can mean that capitula in the exotic range may wilt under the attack levels of R. conicus, leaving all larvae dead (Cullen and Sheppard, 2012). R. conicus in its native range can also be quite heavily attacked by Nosema, a microsporidian parasite (Woodburn and Cullen, 1995; Cullen and Sheppard, 2012). In its exotic range R. conicus has also attracted parasitoids (Zwölfer and Harris, 1984; Wilson and Andres, 1986), but at quite low rates.
Means of Movement and DispersalTop of page
It does not seem that R. conicus has managed to spread to new regions unaided. Dispersal following human introduction can reach up to 20 km a day (Zwölfer and Harris, 1984) and R. conicus can quickly spread over a large area (Hodgson and Rees, 1976). However, dispersal only takes place in spring and in the absence of suitable nearby hosts.
Pathway CausesTop of page
Impact SummaryTop of page
Economic ImpactTop of page
The economic benefits of R. conicus as a biological control agent against the exotic thistle Carduus nutans have been very significant in Canada, the USA and New Zealand. This agent has been less effective in Australia. The size of these economic benefits has not been quantified financially for any of these countries; however, the success of control due to other agents in Australia has been estimated at $69M for whole of life (Page and Lacey, 2006). Based on the relative distribution and abundance of C. nutans in these other countries, the benefits to New Zealand resulting from the impacts of R. conicus are likely to be a similar order of magnitude to Australia, while in North America the benefits are likely to be very much higher, perhaps an order of magnitude higher or more.
Environmental ImpactTop of page
Impact on Biodiversity
R. conicus reduces seed production of the native North American thistle species it is able to attack (Turner et al., 1987; Louda, 1998). This can have a direct impact on population density of the affected species when they are seed limited (Louda et al., 1997). The extent of the impact therefore relates to the degree to which the thistle populations are already threatened by other factors, and the level of damage R. conicus is able to cause, given the proximity and abundance of its normal hosts and the degree of synchrony in flowering between these and the native thistles (Russell et al., 2007).
When present in high densities R. conicus may also impact native insects in the capitula of the native thistles (Tatyana and Louda, 2011). Some native thistle populations may be pushed to extinction by this weevil under some circumstances (Louda and Potvin, 1995).
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Cirsium canescens||National list(s)||USA||Louda et al. (1990)|
|Cirsium ownbeyi||National list(s)||USA||DePrenger-Levin et al. (2010)|
|Cirsium pumilum var. hillii||National list(s)||USA||Sauer and Bradley (2008)|
|Cirsium pitcheri (Pitcher's thistle)||NatureServe; USA ESA listing as threatened species||Illinois; Indiana; Michigan; Wisconsin||Herbivory/grazing/browsing||US Fish and Wildlife Service (2010a)|
|Cirsium vinaceum (Sacramento Mountains thistle)||NatureServe; USA ESA listing as threatened species||New Mexico||Herbivory/grazing/browsing||US Fish and Wildlife Service (2010b)|
Social ImpactTop of page
The effective control of the thistle Carduus nutans will have generated significant social benefits for the farmers this weed was preventing from maintaining a viable livestock business.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Highly mobile locally
- Has high reproductive potential
- Host damage
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Negatively impacts animal/plant collections
- Competition (unspecified)
- Interaction with other invasive species
- Parasitism (incl. parasitoid)
- Difficult to identify/detect as a commodity contaminant
UsesTop of page
The economic and social benefits of R. conicus are due to its effective control of the invasive thistle Carduus nutans, which impacts on the grazing and pastoral industries and farmer livelihoods.
Uses ListTop of page
- Biological control
Similarities to Other Species/ConditionsTop of page
The other four species in the genus Rhinocyllus also feed in the capitula of members of the tribe Cardueae (in the Asteraceae). These are R. oblongus Cap. (found in Spain, Italy, Greece and Turkey), R. depressirostris Boh. (southern Russia), R. remaudieri Hoffm. (Iran) and R. schoenherri Cap. (Caucasus). Of these species, only R. oblongus is sympatric with R. conicus in the latter’s native range (Zwölfer and Harris, 1984), and may indeed be a subspecies (Klein and Seitz, 1994).
A sixth species, R. inquilinus Gyll., described from Finland, appears to be a small individual of R. conicus (Zwölfer and Harris, 1984).
No other species of Rhinocyllus have been introduced as biological agents or are found outside Europe, North Africa and western Asia.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
The movement of R. conicus across USA state borders is now prohibited, an attempt by the authorities to contain the impact of this weevil on native thistles.
Gaps in Knowledge/Research NeedsTop of page
More work is needed to better understand the genetic patterns underlying ecotypic differences in R. conicus and to see if the different ecotypes show any degree of reproductive incompatibility.
ReferencesTop of page
Alonso-Zarazaga MA, Talamelli F, 2011. Fauna Europaea. Fauna Europaea (online). http://www.faunaeur.org/full_results.php?id=248915
Andres LA, Rees NE, 1995. Musk thistle. Biological control in the western United States, 3361 [ed. by Niechols, J. R. \Andres, L. A. \BeardsleyJW, \Geoden, R. D. \Jackson, C. G.]. California, USA: University of California Division of Agriculture and Natural Resources, 248-251
Boldt PE, Kok LT, 1982. Bibliography of Rhinocyllus conicus Froel. (Coleoptera: Curculionidae), an introduced weevil for the biological control of Carduus and Silybum thistles. Bulletin of the Entomological Society of America, 28(4):355-358
Csiki E, 1934. Coleoptera, Curculionidae, Subfamily Hyperinae. In: Schenkling S, ed. Coleopterorum Catalogus. Part 137. Berlin, Germany: Dr W. Junk
Cullen J, Sheppard AW, 2012. Carduus nutans L. - nodding thistle. In: Biological control of weeds in Australia 1960 to 2010 [ed. by Julien, M. \McFadyen, R. \Cullen, J.]. Melbourne, Australia: CSIRO Publishing, 118-130
DePrenger-Levin ME, Grant TA III, Dawson C, 2010. Impacts of the introduced biocontrol agent, Rhinocyllus conicus (Coleoptera: Curculionidae), on the seed production and population dynamics of Cirsium ownbeyi (Asteraceae), a rare, native thistle. Biological Control, 55(2):79-84. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WBP-50KWFY0-1&_user=10&_coverDate=11%2F30%2F2010&_rdoc=2&_fmt=high&_orig=browse&_origin=browse&_zone=rslt_list_item&_srch=doc-info(%23toc%236716%232010%23999449997%232353748%23FLA%23display%23Volume)&_cdi=6716&_sort=d&_docanchor=&_ct=11&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=78efe49241aa2b1324dbea78e84c1945&searchtype=a
Gassmann A, Kok L-T, 2002. Musk thistle (nodding thistle). In: Biological control of invasive plants in the eastern United States [ed. by Driesche, R. Van \Blossey, B. \Hoddle, M. \Lyon, S. \Reardon, R.]. Washington, DC, USA: USDA Forest Service, 229-245
GBIF, 2012. Global Biodiversity Information Facility. Global Biodiversity Information Facility (GBIF). http://data.gbif.org
Goeden RD, 1995. Milk thistle. Biological control in the western United States, 3361 [ed. by Niechols, J. R. \Andres, L. A. \Beardsley, J. W. \Geoden, R. D. \Jackson, C. G.]. California, USA: University of California Division of Agriculture and Natural Resources, 245-247
Goeden RD, Ricker DW, 1985. Seasonal asynchrony of Italian thistle, Carduus pycnocephalus, and the weevil, Rhinocyllus conicus (Coleoptera: Curculionidae), introduced for biological control in southern California. Environmental Entomology, 14(4):433-436
Harris P, 1981. Carduus nutans L., nodding thistle and C. acanthoides L., plumeless thistle (Compositae). Biological control programmes against insects and weeds in Canada 1969-1980 [ed. by Kelleher, J.S.\Hulme, M.A.]. Slough, UK: Commonwealth Agricultural Bureaux, 115-126
Harris P, Zwölfer H, 1971. Biological control of weeds in Canada, 1959-1968. 29. Carduus acanthoides L., welted thistle, and C. nutans L., nodding thistle (Compositae). Technical Communications. Commonwealth Institute of Biological Control, 4:76-9
Jessep CT, 1981. Nodding thistle receptacle weevil, Rhinocyllus conicus (Froelich), life cycle. DSIR Information Series, 105. 37
Klein M, Seitz A, 1994. Geographic differentiation between populations of Rhinocyllus conicus Frölich (Coleoptera: Curculionidae): concordance of allozyme and morphometric analysis. Zoological Journal of the Linnean Society, 110(2):181-191
Laing JE, Heels PR, 1978. Establishment of an introduced weevil Rhinocyllus (Col., Curculionidae) for the biological control of nodding thistle Carduus nutans (Compositae) in Southern Ontario. Proceedings of the Entomological Society of Ontario, 109:3-8
Louda SM, Potvin MA, Collinge SK, 1990. Predispersal seed predation, postdispersal seed predation and competition in the recruitment of seedlings of a native thistle in Sandhills prairie. American Midland Naturalist, 124(1):105-113
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26/03/12 Original text by:
A Sheppard, CSIRO Entomology, Australia
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