The Natural Host Range of Beetle Species Feeding on Broom, Cytisus scoparius (L.) Link (Fabaceae), in Southwest Europe

The natural host range of beetles feeding on broom ( Cytisus scoparius ) and 14 other species (including six other Cytisus species) in the tribe Genisteae was investigated at 39 sites in Spain , Portugal and France in 1989 and 1992 as part of a biological control programme for broom . Data on host - plant associations were analyzed for 36 phytophagous beetle species from 18 sites , and host records were listed for an additional 58 species . Nine species were apparently restricted to the genus Cytisus : Cryptocephalus octoguttatus, Gonioctena olivacea, Bruchidius lividimanus, B. villosus, Exapion elongatissimum, E. fuscirostre, Lepidapion sp . 1 , Polydrusus confluens and Tychius parallelus. These field records suggest a narrower host - plant range for some beetle species than laboratory host - range tests , and may assist in interpreting host - plant associations reported in the literature . Beetle species with a restricted host - plant range were rarely found on related non - host plants .     

Potential non-target impact of Microctonus aethiopoides Loan (Hymenoptera: Braconidae) on Cleopus japonicus Wingelmüller (Coleoptera: Curculionidae), a biocontrol agent for putative release to control the butterfly bush Buddleja davidii Franchet in New Zealand

Abstract  Cleopus japonicus Wingelmüller (Coleoptera: Curculionidae) is being considered for release to control buddleia Buddleja davidii in New Zealand. As part of the pre-release testing, Moroccan and Irish biotypes of the solitary endoparasitoid Microctonus aethiopoides Loan (Hymenoptera: Braconidae) were evaluated for potential non-target impacts on adult C. japonicus should release occur. Laboratory experiments evaluated both the behavioural and physiological suitability of C. japonicus to both biotypes of the parasitoid. Parasitoid behavioural attraction was assessed using the pathenogenic bacterium Serratia marcescens (Enterobactereaceae), as an indicator of ovipositor penetration. Physiological suitability was assessed by comparing parasitism of C. japonicus with the natural hosts of the respective parasitoid biotypes. The parasitoid-bacteria study showed that C. japonicus was behaviourally acceptable to both Moroccan and Irish M. aethiopoides , with the two experiments producing 34% and 8% mortality, respectively. Cleopus japonicus did not support development of either Moroccan or Irish M. aethiopoides biotypes. None of the weevils dissected at the end of the experiment contained immature parasitoids. Comparison between unexposed and parasitoid-exposed C. japonicus found no difference in premature mortality during the experiment nor in the number of fully reproductive females at its conclusion. The results of this study predict that should C. japonicus be released, the potential impact of M. aethiopoides on field populations will be negligible.     

Studies on the feeding of Sitona lineatus L. (Coleoptera: Curculionidae) on white clover (Trifolium repens L.) seedlings

Measurements of feeding damage by sitona weevil (Sitona lineatus L.) adults on differing numbers of seedlings of white clover (Trifolium repens L.) at the first and fourth trifoliate leaf stage were made in the glasshouse at 20°C. S. lineatus consumed more of the trifoliate component of the seedling. Sitona adults caused significant yield reduction at all levels of plant population. Total clover consumption increased with increasing size of sitona population, but consumption per adult weevil was reduced.     

Host specificity testing and suitability of a European biotype of the braconid parasitoid Microctonus aethiopoides as a biological control agent against Sitona lepidus (Coleoptera: Curculionidae) in New Zealand

The European biotype of the parasitoid Microctonus aethiopoides Loan (Hymenoptera: Braconidae) is being considered for release against Sitona lepidus Gyllenhal (Coleoptera: Curculionidae) in New Zealand. Host specificity was evaluated in the laboratory using both endemic and introduced weed biological control curculionid species, with 12 no-choice and three choice experiments carried out comparing the S. lepidus and test weevils. Two further no-choice tests used the Moroccan M. aethiopoides biotype to compare attack rate between European and Moroccan M. aethiopoides, the latter released in 1982 to control the lucerne pest S. discoideus. Across all experiments, total parasitism of S. lepidus was 69% compared with 15% for the test weevils. European M. aethiopoides was able to develop in the native weevils Irenimus aequalis, Nicaeana cervina, Catoptes cuspidatus, Protolobus porculus and Steriphus variabilis with parasitism rates of 13, 28, 2, 7 and 8%, respectively. These levels were significantly less than those in the corresponding S. lepidus control. Total parasitism of I. aequalis and C. cuspidatus increased significantly in the presence of S. lepidus than recorded under no-choice conditions. The presence of European M. aethiopoides caused minor, if any, test weevil mortality prior to the onset of prepupal emergence and there was no significant reproductive suppression in parasitoid-exposed test weevils. Parasitism of the introduced weed control agent R. conicus by European M. aethiopoides was significantly lower (1.1%) compared to the Moroccan biotype (47.5%). Based on these and other experiments, should the European M. aethiopoides be released as a biological control agent of S. lepidus, its ecological impacts are likely to be less severe than those already exhibited by the Moroccan M. aethiopoides.     

Sitona weevils (Coleoptera: Curculionidae) as agents for rapid transfer of nitrogen from white clover (Trifolium repens L.) to perennial ryegrass (Lolium perenne L.)

The effects of root feeding by larvae of Sitona hispidulus (F.) (a common weevil pest of white clover) on the rate of transfer of nitrogen between plants of white clover (Trifolium repens L.) and perennial ryegrass (Lolium perenne L.) were investigated using a nutrient slant board technique. Clover plants, labelled with ¹⁵N were grown adjacent to ryegrass plants and were either infested with Sitona larvae or not infested. Ryegrass plants associated with the infested clover plants had a significantly higher dry matter yield and nitrogen content (75% and 74% respectively) than the uninvested plants, after 33 days exposure to insect herbivory. It was concluded that root feeding insects could play an important role in the cycling of nitrogen in grass/clover swards.     

Demography of the invasive shrub Scotch broom (Cytisus scoparius) at Barrington Tops,New South Wales: insights for management.

The exotic shrub Scotch Broom (Cytisus scoparius) has invaded large areas of eucalypt woodland at Barrington Tops, New South Wales, where it forms dense stands that have significant impacts on vegetation structure, flora and fauna. Data are presented from four 25 m² plots, which have been studied since 1985. Two plots were located in uniform broom thickets of different ages, and two were located across the margins of broom stands, which have since expanded to cover the entire plots. All broom plants in the plots (other than young seedlings, which were counted) were mapped, tagged and monitored annually. New seedlings appeared annually, but there was no relationship between their numbers (varying between years) and subsequent recruitment of older plants. The probability of seedlings reaching first flowering was less than 2%, and of surviving to mature size (>10 cm² basal area) was negligible. Seedlings mainly died through suppression (shade). Individuals less than 50 cm high were also browsed. Recruitment occurred only where light levels were high, either before closure of the broom canopy or after senescence had led to canopy opening. From approximately 12–30 years after initial invasion, broom stands underwent self‐thinning of mature plants, accelerated by collapse of plants on to each other. Recruitment of new, maturing plants, after this period produced a stand that was less dense than that found after initial invasion. Broom is creating more disturbance‐prone environments due to its impacts on other biota, likely alterations to the fire regime, and by harbouring feral pigs. Further disturbance favours broom, and elsewhere it has resulted in massive seedling regeneration. While fire or other disturbance can be used to stimulate germination, and thereby reduce a large part of the soil seed bank, denser broom infestations are likely to result unless follow‐up treatments can be applied over long time periods. A wiser management option, at least in the short term, may be avoidance of all disturbance, especially for stands of mature broom.     

Evaluation of Cyphocleonus trisulcatus (Coleoptera: Curculionidae) as a potential biological control agent for Leucanthemum vulgare in North America

Leucanthemum vulgare, a perennial forb native to Eurasia, has become invasive in several other parts of the world. Since there is a lack of methods suitable for sustainable management of this plant across invaded landscapes, a biological control project has been initiated in North America and Australia. We evaluated the potential of the root-feeding weevil Cyphocleonus trisulcatus as a biological control agent for L. vulgare in North America by investigating its impact on L. vulgare and studying its larval host range under no-choice, multiple-choice cage and open-field conditions. Our impact experiment revealed that larval feeding by C. trisulcatus can severely damage and even kill plants. No-choice tests conducted with 41 non-target species showed that C. trisulcatus can develop on five closely related species. In a multiple-choice cage test established with three of these species, the medicinal plant Matricaria chamomilla and the native North American M. occidentalis were also attacked, but to a lower degree than L. vulgare. An open-field test conducted with four varieties of the ornamental Shasta daisy (Leucanthemum × superbum and L. × maximum) revealed similar attack on Shasta daisy as on L. vulgare. Due to the popularity of Shasta daisies, we suggest that C. trisulcatus, despite its potential to suppress L. vulgare populations, is not a suitable biological control agent for North America. However, C. trisulcatus may still be a potential biological control agent for L. vulgare in regions where Shasta daisy is less popular, such as Australia. Further host-range tests with species native to Australia as well as with additional species important for the horticultural industry in Australia are currently under way.     

Gallerucida bifasciata (Coleoptera: Chrysomelidae), a potential biological control agent for Japanese knotweed (Fallopia japonica)

Host specificity of foreign natural enemies are becoming more and more critical in classical biological control programs, as concerns about potential risk from introduced biocontrol agents have been increasing recently. Understanding the insect's fundamental and ecological host ranges is the first step in determining the potential for introduction of an insect to control invasive plants. Japanese knotweed, Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is an invasive weed in the United States and Europe. A leaf beetle, Gallerucida bifasciata (Coleoptera: Chrysomelidae) is an important natural enemy attacking this plant in Asia. However, its host range records were ambiguous. This study examined the beetle's host specificity through a set of choice and no-choice tests in the laboratory and field in its native China. Gallerucida bifasciata larvae were able to complete development on seven of 87 plant species in larval development tests, while adults fed and oviposited on 10 plants in no-choice tests. Multiple choice tests showed adults strongly preferred Fallopia japonica, Persicaria perfoliata (L.) H. Gross and Polygonum multiflorum Thunb over all other plants. Open field tests and field surveys further revealed that these three species were in its ecological host range. The results of this study suggest that G. bifasciata is a potential promising agent for control of Japanese knotweed in the United States and Europe, although additional host specificity tests and risk assessment should be completed.     

Distribution and abundance of three species of Sitona (Coleoptera: Curculionidae) in grassland in England

The distribution and abundance of weevils of the genus Sitona was assessed at 16 sites in England. The sites were chosen to cover a wide range of geographical and agricultural regions. Adult weevils were sampled on four occasions, from May to August 1993. Numbers of adults, larvae and pupae were determined for each site on four occasions from June to September 1994. Numbers of larvae declined throughout the year in which they were sampled and pupae were most numerous in July. The population density of adults increased through the sampling period in both years. Sites close to pulse crops had the largest population levels of weevils, with Sitona lineatus L. the dominant species. No differences in the population levels of Sitona hispidulus (F.) or S. flauescens (Marsh.) were detected. Evidently populations of the latter two species are fairly constant in grassland and most of the variation in the population densities of these three species found in English grassland can be attributed to the migratory behaviour of S. lineatus and the proximity of pulse crops.     

First record of Sitona discoideus Gyllenhal 1834 (Coleoptera: Curculionidae) on Norfolk Island

Abstract

The first records of Sitona discoideus and Trifolium repens on Norfolk Island are reported. The identities of both species were confirmed using morphological criteria and nucleotide sequences. Sequence data from the mitochondrial gene cytochrome oxidase subunit 1 of the Norfolk Island S. discoideus specimens were compared with 33 specimens from Australia, New Zealand and France. These data represent the first published sequences for S. discoideus, and suggest that Australia or New Zealand was the source of Norfolk Island's population. The implications of the introduction of S. discoideus to Norfolk Island are discussed.     

Biology and prey range of Cryptognatha nodiceps (Coleoptera: Coccinellidae), a potential biological control agent for the coconut scale, Aspidiotus destructor (Hemiptera: Diaspididae)

Cryptognatha nodiceps Marshall is an important natural enemy used in biological control programmes against Aspidiotus destructor (Signoret), an introduced pest of coconut in many countries. In order to increase our understanding of C. nodiceps, the present study to quantify aspects of the developmental, reproductive and feeding biology under constant environmental conditions (26±2°C, 55-65% RH) was carried out. In addition, methods for culturing the scale and the beetle were developed. The average duration of development was 19.2±0.1 days for males and 19.6±0.2 days for females. Female longevity and lifetime fecundity was 35.6±5.1 days and 141.0±35.6 eggs, respectively. Life table statistics were as follows: reproductive rate, 9.99, intrinsic rate of increase, 0.09, finite rate of increase, 1.1, doubling time, 7.8 days and generation time, 41.1 days. Studies on feeding biology showed that C. nodiceps was oligophagous. The coccinellid fed and reproduced on prey species in two families, Diaspididae and Coccidae. Adults also fed on the coconut whitefly Aleurodicus cocois (Curtis) (Aleyrodidae) but no reproduction occurred on this prey. Aphididae, Psyllidae and Pseudococcidae were not fed upon.     

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

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

Rapid cold-hardening response in Ophraella communa LeSage (Coleoptera: Chrysomelidae), a biological control agent of Ambrosia artemisiifolia L.

In China, an unintentionally introduced beetle Ophraella communa shows good control efficiency against invasive common ragweed, Ambrosia artemisiifolia. With the aim of understanding the capacity of O. communa to overcome sub-zero low temperature rapidly, the longevity, survival and fecundity of the beetle adults were studied at conditions of different rapid cold-hardening in the laboratory. The results showed that approximately 20% female or male individuals still survive at ?12°C for 2 h, thus the discriminating temperature for rapid cold-hardening of both female and male beetles is ?12°C. The survival, longevity and fecundity of adult beetles were significantly affected by rapid cold-hardening treatments. Compared to the control, the survival rates of adult beetles experiencing 2 h of rapid cold-hardening at 2, 5 and 8°C prior to exposure to its discriminating temperature (?12°C) were significantly increased. Adult longevity and fecundity were significantly higher after pre-treatment at 2, 5 and 8°C for 2 h than those of the control. Both longest longevity and maximum fecundity of adult beetles were observed at the rapid cold-hardening of 2°C. Our experiment implies that the cold-tolerance ability of O. communa can be rapidly increased after rapid cold-hardening treatments, which reveals an important ecological significance for population establishment and expansion of O. communa in the invaded areas of common ragweed.     

Biology and host specificity of Aulacobaris fallax (Coleoptera: Curculionidae), a potential biological control agent for dyer’s woad,Isatis tinctoria (Brassicaceae) in North America

Dyer’s woad, Isatis tinctoria, a plant of Eurasian origin is a problematic weed in western North America against which a classical biological weed control programme was initiated in 2004. Three European insect species were selected as candidate agents to control this invasive species, including the root‐mining weevil Aulacobaris fallax. To determine its suitability as an agent, the biology and host specificity of A. fallax were studied in outdoor plots and in the field between 2004 and 2006 in its native European range. Aulacobaris fallax is a univoltine species that lays its eggs from March to August into leaf stalks and roots of dyer’s woad. Larvae mine and pupate in the roots and adults emerge from August to October. Up to 62% of the dyer’s woad plants at the field sites investigated were attacked by this weevil. In no‐choice host‐specificity tests, A. fallax attacked 16 out of 39 species and varieties within the Family Brassicaceae. Twelve of these are native to North America. In subsequent multiple‐choice tests, seven species, all native to North America, suffered a similar level of attack as dyer’s woad, while none of the European species were attacked. Our results demonstrate the importance of including test plant species that have not co‐evolved with the respective candidate agent. In sum, we conclude that the risk of non‐target effects is too high for A. fallax to be considered as a biological control agent for dyer’s woad in the United States.     

Revised methods for the mass-rearing of the spotted knapweed biological control agent,Cyphocleonus achates (Coleoptera: Curculionidae), in field corrals

Cyphocleonus achates (Fahraeus), a root-feeding weevil introduced from Eurasia, is an effective biological control agent against spotted knapweed, Centaurea stoebe L. ssp. micranthos. Because C. achates is univoltine and does not fly, distribution of the weevil has been slow. To hasten the weevil's distribution, a rearing effort using field corrals was initiated at a facility in Corvallis, Montana. Procedures for mass-rearing the weevil in field corrals are described, with an emphasis on improvements over earlier methods. The described field-corral approach is effective and appropriate for producing C. achates for distribution in the western United States.     

A prospective model for the phenology of Microctonus hyperodae (Hymenoptera: Braconidae), a potential biological control agent of argentine stem weevil in New Zealand

A predictive phenological model is described for the parasitoid Microctonus hyperodae, introduced to New Zealand as a potential biological control agent against Argentine stem weevil Listronotus bonariensis. The model is based on development/temperature relationships obtained from experiments on the parasitoid in quarantine prior to its release, allowing early predictions of its phenology in different parts of the target pest's New Zealand range. In particular the model was used to predict the number of parasitoid generations each year, the degree of temporal synchrony between parasitoid adults and the susceptible adult pest stage, the order of parasitism and reproduction in the pest's life cycle as a possible basis for a simplified, discrete host/parasitoid population model, and the likely significance of ecotypic differences in development and diapause characteristics of the parasitoid. These applications demonstrate the potential for simple models to help in climate matching of classical biological control agents and estimation of their interaction with pest dynamics, using data obtainable prior to their introduction and release. In addition the model proved useful as a decision aid during the release programme, by indicating the likely effects of unusual weather and the need or otherwise for further parasitoid releases.     

Laboratory host range of Austromusotima camptozonale (Lepidoptera: Crambidae), a potential biological control agent of Old World climbing fern,Lygodium microphyllum (Lygodiaceae)

Old World climbing fern, Lygodium microphyllum, is one of the most serious invasive weeds impacting south Florida and development of biological control is crucial for sustainable management. Larvae of a small moth, Austromusotima camptozonale, were discovered defoliating L. microphyllum in Australia. Preliminary testing suggested this moth was a Lygodium specialist. Laboratory host range testing was conducted on 65 species of test plants, from 31 families, comprising seven Lygodium species, four close relatives, 45 other species of ferns and fern allies, eight agricultural crops and one gymnosperm species plus the primary host L. microphyllum. Significant oviposition occurred only on other species of Lygodium. No eggs were laid on the agricultural crops, or about half the species of non-Lygodium ferns and fern allies tested. Oviposition on the other non-Lygodium ferns was very low, except on Anemia adiantifolia and Blechnum serrulatum, which received modest egg loads, but did not support development to adult. Larval feeding was low to non-existent on all the non-Lygodium species. Larvae developed to adult only on the native, American climbing fern L. palmatum, and to a lesser extent on L. japonicum. Lygodium japonicum is a naturalized invasive weed in the United States. Colonies of A. camptozonale were unable to persist on L. palmatum and died out in two to seven generations. Freezing winter temperatures in states where L. palmatum occurs would be lethal to A. camptozonale. It was concluded that A. camptozonale would pose no threat to native or cultivated plants in North America or the Caribbean and should be considered as a weed biological control agent against L. microphyllum.     

Post-establishment assessment of host plant specificity of Arytainilla spartiophila (Hemiptera: Psyllidae), an adventive biological control agent of Scotch broom,Cytisus scoparius

Scotch broom, Cytisus scoparius (Fabaceae), is a shrub native to Europe that is invasive in the USA, New Zealand and Australia. The psyllid Arytainilla spartiophila has been purposely introduced to Australia and New Zealand as a biological control agent of C. scoparius, but is an accidental introduction to California. Lupines (Lupinus spp.) are the closest native taxon to Cytisus in North America, and are therefore considered to be at the highest risk for non-target damage. However, because no lupines are native to Australia or New Zealand, only one imported forage species was evaluated during prior host specificity testing. We conducted a laboratory nymphal transfer experiment, a field choice experiment and a field survey to assess risk to three lupine species (Lupinus albifrons, Lupinus bicolor and Lupinus formosus). In the laboratory, 20% of third-instar nymphs were able to develop to adulthood on L. formosus but not on the other lupine species, while 40% completed development on C. scoparius. In the field experiment, potted lupine and C. scoparius plants were placed beside large infested C. scoparius plants; oviposition occurred on all the potted C. scoparius plants, but on none of the lupines. In the field survey, no A. spartiophila eggs or nymphs were found on naturally occurring lupines growing adjacent to infested C. scoparius. The results indicate that A. spartiophila is not likely to damage or reproduce on lupines in the field. This study provides an example of how field studies can help clarify the host specificity of biological control agents.     

Host range of Ceutorhynchus assimilis (Coleoptera: Curculionidae), a candidate for biological control of Lepidium draba (Brassicaceae) in the USA

Ceutorhynchus assimilis has been selected as a potential biological control agent of Lepidium draba, which is a Eurasian invasive weed in North America. Preliminary studies indicated specificity of this weevil collected in southern France on L. draba. This result was in discord with the pest status of C. assimilis found in the literature. Host-specificity tests based both on field and laboratory experiments showed heterogeneity in the host spectrum of the weevils reared from different host-plants as determined by larval development. However, no distinguishable morphological differences could be visually detected between the populations feeding on different host-plants. All sampled populations of weevils were polyphagous as adults. Weevils reared from L. draba were specific to this plant for their complete larval development. Conversely, populations living on other wild and cultivated Brassicaceae species were not able to use L. draba as a host plant. Such differentiation is further highlighted by other biological aspects such as plant infestation rates, sex-ratio, duration of larval development, and differences in the timing of their life cycles. These results demonstrate that C. assimilis, an insect species formerly considered as a pest of Brassicaceae, is characterized by its host-range variability, with one population being potentially useful in the biological control of L. draba. Moreover, this example points to the need to test multiple populations of biological control agents in assessing risk.     

Eustenopus villosus (Coleoptera: Curculionidae) Feeding of Herbicide-Resistant Yellow Starthistle (Centaurea solstitialis L.)

A population of yellow starthistle (Centaurea solstitialis L.) near Dayton, Washington developed herbicide resistance in response to repeated applications of picloram and other auxin-type herbicides. Laboratory and field experiments were conducted in 1998 to determine host acceptability and suitability of this herbicide-resistant yellow starthistle population to the biological control weevil Eustenopus villosus (Boheman) (Coleoptera: Curculionidae). In choice and no-choice feeding and oviposition experiments using excised buds, the weevil did not demonstrate a consistent preference for either herbicide-resistant (R) or -susceptible (S) yellow starthistle. When caged on buds of intact plants, the E. villosus feeding rate of 97% did not differ between R and S types. Host plant suitability, measured as larval damage and development to adult weevils, was equivalent in R and S types, with weevils maturing in 46% of the R and in 32% of the S capitula bearing oviposition scars. The number of viable achenes per capitulum was reduced by 87% due to larval feeding, with no difference between R and S types. Observations at the field site where resistance was found revealed oviposition scars on 78% of the late-bud-stage capitula on 23 June 1998 and 73% of the flowering and postflowering capitula on 15 August 1998. Selection for herbicide resistance has not created host incompatibility for E. villosus nor reduced the effectiveness of E. villosus as a biological control agent.