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.     

Assessment of risk of attack to safflower by Ceratapion basicorne (Coleoptera: Apionidae), a prospective biological control agent of Centaurea solstitialis (Asteraceae)

Ceratapion basicorne (Coleoptera: Apionidae) is a prospective biological control agent of yellow starthistle (Centaurea solstitialis, Asteraceae: Cardueae), which is an important invasive alien weed in the western United States. Previous studies have shown that it is possible for this insect to oviposit on and complete development on safflower (Carthamus tinctorius) under no-choice laboratory conditions; however, it has never been reported as a pest of safflower. Field experiments were conducted at three sites in eastern Turkey during 3 years to evaluate the risk of attack on safflower by this insect in its native range. At two sites where C. basicorne was the only apionid observed, no safflower plants were attacked despite high attack rates on yellow starthistle test plants (48–98% of plants infested). At a third site, where C. basicorne and three other species in the same genus; C. scalptum, C. orientale, and C. onopordi were present, 8–26% of safflower plants were infested, but none of the insects reared from safflower during 3 years were C. basicorne. Other authors have reported rearing C. basicorne from field-collected plants of only Ce. solstitialis, Ce. cyanus, Ce. depressa, and Cnicus benedictus. Our results indicate that C. basicorne does not attack safflower under field conditions and that its introduction would not pose a risk to this crop.     

Assessing the biological control of yellow starthistle (Centaurea solstitialis L): prospective analysis of the impact of the rosette weevil (Ceratapion basicorne (Illiger))

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Host range and risk assessment of Zygogramma bicolorata,a defoliating agent released in South Africa for the biological control of Parthenium hysterophorus

Parthenium hysterophorus (parthenium) is a weed of international importance and is spreading rapidly in sub-Saharan Africa. Consequently, it has been targeted for biocontrol in South Africa since 2003. Based on precedents elsewhere in the world, the defoliating beetle Zygogramma bicolorata was prioritised as a candidate agent. Although no-choice tests, involving some 48 test plant species, indicated a significant preference for parthenium, significantly reduced feeding and oviposition was recorded on some species. Multiple-choice tests resolved many of these non-target results; however, Helianthus annuus (sunflower) was still selected for oviposition and feeding. Of the 12 sunflower cultivars tested, four were selected for oviposition, while two were selected for oviposition and feeding. These six cultivars were then subjected to larval development trials, together with three native and two weed species (in the Asteraceae). These trials showed high levels of complete development on parthenium, significantly reduced development on sunflower cultivars, and partial development on only one of the weed species. Finally, a risk assessment was conducted on the six sunflower cultivars to quantify Z. bicolorata feeding and reproductive performance. Feeding risk calculations revealed these cultivars to have an extremely low risk (<0.2%) of supporting Z. bicolorata feeding and development. Similarly, reproductive risk calculations showed a very low risk (<0.16%) of supporting viable Z. bicolorata populations. These data are supported by findings from both the native (Mexico) and introduced ranges (Australia, India) of Z. bicolorata, where it has never been recorded as a pest of sunflower. These considerations were accepted by the regulatory authorities and in August 2013, Z. bicolorata became the second insect agent to be released in South Africa for the biocontrol of parthenium.     

Host specificity ofPareuchaetes insulata (Lep.: Arctiidae), a biological control agent forChromolaena odorata (Compositae)

Larvae of the arctiid moth,Pareuchaetes insulata, from Florida fed on the leaves ofChromolaena odorata, a serious composite, alien weed in Natal, South Africa. In starvation test trials using 48 plant species,P. insulata completed its development onC. odorata andAgeratum houstonianum. Subsequent attempts to cultureP. insulata on these two plants was only successful onC. odorata. The biology ofP. insulata is similar to that of two other well studiedPareuchaetes species namelyP. pseudoinsulata andP. aurata aurata. Repeated defoliation ofC. odorata byP. insulata could contribute to its control as has been found withP. pseudoinsulata in Guam.Pareuchaetes insulata has been approved for release as a biological control agent ofC. odorata in South Africa.     

Host range of the European,rhizome-stem feeding scale Rhizaspidiotus donacis (Hemiptera: Diaspididae), a candidate biological control agent for giant reed,Arundo donax (Poales: Poaceae) in North America

The armored scale Rhizaspidiotus donacis (Leornardi) was evaluated as a potential biological control agent of the invasive reed grass Arundo donax in North America. No-choice tests, native range field surveys and non-target host exposures were used to determine the fundamental host range of the scale collected from Caloma, Spain and Perpignan, France. Thirty-five species, including two genotypes of A. donax and seven ecotypes of Phragmites australis, along with closely related grasses, economic grasses and habitat associates were tested. In quarantine no-choice testing using releases of 200 crawlers per plant, normal development of R. donacis was observed on A. donax and A. formosana, with very limited survival to the adult stage on Spartina alterniflora and Leptochloa spp. In follow-up studies using 1000 crawlers per plant, 10 live adult females were found on Leptochloa virgata, and one adult female on Spartina alterniflora, but average adult female abundance per plant was (2580%) 26-times lower on L. virgata and over (39,090%) 100-times lower on S. alterniflora than on A. donax. Field surveys were conducted at five locations in Spain and France at which A. donax infested with R. donacis, co-occurred with two non-target species of concern and R. donacis was only found on A. donax. Six-month field host exposures in Spain using potted Leptochloa plants entwined with heavily infested A. donax confirmed that R. donacis is specific to Arundo under field conditions. Based on our results, the scale R. donacis appears to be specific to the genus Arundo and is unlikely to harm native or cultivated plants in the Americas.     

Laboratory studies on the biology and host range of Dichrorampha odorata (Lepidoptera: Tortricidae), a biological control agent for Chromolaena odorata (Asteraceae)

Dichrorampha odorata (Lepidoptera: Tortricidae) is a moth from Jamaica whose larvae bore into, and kill, the shoot tips of the invasive alien plant, Chromolaena odorata (L.) King and Robinson (Asteraceae). This study reports aspects of the biology of D. odorata, and also determined the host specificity (larval and adult no-choice trials) of the moth. Adults were short lived (ranging from 2 to 7 days), with females laying a mean of 15.4 eggs. Eggs took 9 days to hatch, larvae 20–23 days to develop and the pupal stage lasted 11–12 days, giving an overall lifecycle period of 41–45 days. Larval no-choice tests using 34 asteraceous test species indicated that only C. odorata could sustain complete development of D. odorata to adulthood, although there was slight initial boring 14 test species (plus chromolaena). Results from the adult nochoice trials, in which seven test-plant species were exposed to D. odorata, were consistent with those from larval trials, with larval damage, pupae and adults of D. odorata recorded from only C. odorata. This confirmed that only C. odorata is a suitable host for D. odorata in South Africa. Permission has subsequently been granted for the release of D. odorata in South Africa, thus making it the first shoot-tip attacking agent to be released against C. odorata. It is hoped that in the field, high levels of damage by the moth will reduce the height and therefore competitiveness of C. odorata, thereby contributing to the success of biological control of this plant.     

Host specificity and biology of the weevilNeohydronomus affinis [Coleoptera: Curculionidae] a biological control agent ofPistia stratiotes

Waterlettuce,Pistia stratiotes L., has been successfully controlled in Australia with the weevil,Neohydronomus affinis Hustache. In this study, the weevil was tested in quarantine against 34 plant species in 27 families. In non-replicated no-choice tests, it fed and oviposited on 3 species of duckweeds [(Lemna minor L.,Spirodela punctata (Meyer) Thomps.,S. polyrhiza (L.) Schleid.], frogbit [(Limnobium spongia (Bosc.) Steud.], mosquitofern (Azolla caroliniana Willd) and waterfern (Salvinia mimima Baker). Feeding also occurred on golden club (Orontium aquaticum L.) and one egg was laid on the terrestrial panda plant (Kalanchoe tomentosa Baker). Plants oviposited and fed upon were retested in choice tests. The weevils then fed and oviposited exclusively on waterlettuce. Various aspects of weevil biology, including larval feeding and mining, plant response to weevil attack, and adult behavior are discussed. Florida Agric. Expt. J. series No 8580.     

Release and distribution of Lilioceris cheni (Coleoptera: Chrysomelidae), a biological control agent of air potato (Dioscorea bulbifera: Dioscoreaceae), in Florida

From 2012 to 2015, 429,668 Lilioceris cheni Gressit and Kimoto (Coleoptera: Chrysomelidae) were released in Florida for biological control of air potato Dioscorea bulbifera L. (Dioscoreaceae). In the fall of 2015, a state-wide survey was conducted at 113 randomly selected air potato infestations in order to determine the spatial distribution of L. cheni. Damage due to L. cheni was found at 86% of locations and was low in the Florida panhandle where air potato was relatively uncommon and fewer beetles had been released, and in far south Florida, despite high numbers of beetle releases. On average, beetles travelled 9.5?km from the nearest release site to survey sites from the date of release to the time of the survey, with a maximum distance of nearly 67?km. The rate of spread was estimated at 8.2?km/year under the assumption that beetles present at survey sites migrated or were moved from the nearest release site. Air potato vines produced fewer aerial tubers, the vegetative propagule of air potato, as foliar damage due to L. cheni increased. The results suggest that future research efforts should focus on determining the biotic and abiotic factors that may be limiting establishment in some areas of Florida.     

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.     

Host specificity of <Emphasis Type="Italic">Tectococcus ovatus</Emphasis> (Hemiptera: Eriococcidae), a potential biological control agent of the invasive strawberry guava, <Emphasis Type="Italic">Psidium cattleianum</Emphasis> (Myrtales: Myrtaceae), in Florida

Strawberry guava, Psidium cattleianum Sabine, is a woody tree or shrub native to coastal southeastern Brazil. Strawberry guava was introduced into Florida in the late 1800s as an ornamental species. The plant escaped cultivation and is invading natural areas throughout the southern half of the state. In addition to negative effects on Florida’s native ecosystems, strawberry guava also is a preferred host of the Caribbean fruit fly, Anastrepha suspensa Loew (Diptera: Tephritidae). In total, 57 plant species representing 21 families were included in the host range tests. First instar nymphs of Tectococcus ovatus Hempel fed on two closely related guava species, Brazilian guava (Psidium friedrichsthalianum O. Berg), and Costa Rican guava (Psidium guineense Sw.). However, none of the nymphs completed their development on these two non-target species. The results of the host specificity tests suggest that T. ovatus is a suitable candidate for classical biological control of strawberry guava in Florida.     

Preliminary host specificity testing of Endophyllum osteospermi (Uredinales, Pucciniaceae), a biological control agent against Chrysanthemoides monilifera ssp. monilifera

Chrysanthemoides monilifera ssp. monilifera, indigenous to the Western Cape Province of South Africa, is a serious invader of native vegetation in south-eastern Australia. The rust fungus Endophyllum osteospermi causes witches' brooms on C. monilifera ssp. monilifera in South Africa, and is associated with a reduction in growth and seed production of its host under natural conditions, as well as mortality of severely infected bushes. This rust fungus is considered to be a potential biological control agent for use against C. monilifera ssp. monilifera in Australia. Endophyllum osteospermi has a long latent period, typically between 6 and 24 months between infection and the initiation of witches' brooms. This long latent period makes the logistics of doing traditional host specificity testing, in which all test plant species are inoculated and observed for symptom development, unfeasible for this rust fungus. Germination of aecidioid teliospores and penetration by basidiospores were observed on the surface of excised leaves of 32 test plant species at 4 days after inoculation, and compared to that on C. monilifera ssp. monilifera. Germinating aecidioid teliospores aborted on 14 test plant species, whilst no penetration was attempted on a further 12 test plant species. Penetration only occurred on nine of the 32 test plant species, in addition to C. monilifera ssp. monilifera. Inoculating whole plants of nine selected test plant species confirmed the above results. Therefore, only the test plant species in which penetration occurred, or at least was attempted, need to undergo comprehensive host specificity testing. Pending these results, E. osteospermi may be suitable for release in Australia for the biological control of C. monilifera ssp. monilifera.     

Effect of elevated CO2 on tropical soda apple and its biological control agent Gratiana boliviana (Coleoptera: Chrysomelidae)

Rising atmospheric CO₂ levels could have drastic effects on the performance of invasive weeds and their insect herbivores. Despite the importance of biological control as an effective management tool for environmental weeds, there have been few studies on the potential impact of climate change on the future efficacy of biological control. The objective of the present study was to evaluate the effect of elevated CO₂ on the performance of tropical soda apple Solanum viarum (Solanaceae) and its biological control agent Gratiana boliviana (Coleoptera: Chrysomelidae). We established three levels of CO₂ in environmental growth chambers: ambient (400 ppm), medium (580 ppm) and high (780 ppm). Plants growing at the high level had greater biomass compared to those growing at the ambient and medium levels of CO₂. Leaf water content and the amount of leaf nitrogen were reduced at high compared to ambient or medium CO₂ levels. G. boliviana immature survival and developmental time were negatively affected at high CO₂ but not at medium or ambient levels. Adults were lighter and smaller when reared at the high CO₂ level compared to ambient and medium treatments, while adult fecundity was higher at the medium CO₂ level. Leaf area consumed by fifth instars was lower when feeding on plants grown at the high CO₂ level either inside a Petri dish or on potted plants. These results suggest that beetle performance may be diminished under future climate. However, further studies should incorporate other factors such as temperature and precipitation as well as the evolutionary potential of herbivores and plants to adapt to a changing climate.     

Host specificity testing,release and establishment of Urophora solstitialis (L.) (Diptera: Tephritidae), a potential biological control agent for Carduus nutans L., in Australia

The seed‐fly Urophora solstitialis was imported into Australia from southern France. Previous workers had shown that this seed‐fly limited its attack to selected members of the genus Carduus. Host specificity studies revealed that no native Australian plant tested from the family Asteraceae was a suitable host plant. Permission to release the fly was granted, and it was released in three regions of the New South Wales tablelands. The fly underwent a second generation at two of the release sites, and reasons for its failure to do so at the third are suggested. It has successfully established itself in all three regions, having completed a full seasonal cycle in the field.     

Risk assessment of Episimus unguiculus (Lepidoptera: Tortricidae), a biological control agent of Schinus terebinthifolia (Sapindales: Anacardiaceae) in Hawaii,USA

Schinus terebinthifolia Raddi (Anacardiaceae) is an introduced ornamental tree from South America that has become one of the most invasive weeds in Hawaii and Florida, USA. Exploratory surveys in the plant’s native range from 1950 to 2014 identified several potential biological control agents. One of these is the leaflet rolling moth Episimus unguiculus Clarke (Lepidoptera: Tortricidae), previously known as Episimus utilis Zimmerman. This biological control agent was released in Hawaii in the 1950s where high densities were occasionally observed, leading to partial control of S. terebinthifolia by the 1960s. Larvae are leaf tiers capable of completely defoliating small plants. In order to investigate the release of E. unguiculus in the continental USA, a series of laboratory no- choice, and multiple-choice tests were conducted in Florida, and a preliminary open field test with a native plant in Hawaii. Under the confined laboratory conditions imposed during the no-choice tests, E. unguiculus accepted the economically important Pistacia spp. and several other non-target plants for oviposition and development. However, in the multiple-choice tests E. unguiculus exhibited a clear preference for S. terebinthifolia relative to non-target plants accepted in the no-choice tests. Overall, the results of field observations during surveys in South America and Hawaii and host range studies completed in Hawaii and Florida showed that E. unguiculus is a narrow specialist on S. terebinthifolia, its natural host plant.     

Host specificity of Argopistes tsekooni (Coleoptera: Chrysomelidae), a potential biological control agent of Chinese privet

Chinese privet, Ligustrum sinense Lour., is a perennial semi-evergreen shrub that is a serious invasive weed in the United States. Classical biological control offers the best hope for controlling it in an economic, effective, and persistent way. Host specificity of one of the most promising biological control agents of Chinese privet, a flea beetle, Argopistes tsekooni Chen (Coleoptera: Chrysomelidae), was evaluated in China by using laboratory no-choice and choice tests on 13 species of Oleaceae and eight species in other families that have important economic value. In adult no-choice survival and oviposition tests, the flea beetle fed and survived for 30 d on Syringa oblata Lindl., Jasminum nudiflorum Lindl., and three species in the genus Ligustrum. Females also oviposited on these species, but only larvae from eggs laid on S. oblata and Ligustrum spp. developed successfully. In addition, the beetles did not feed or oviposit on the species of economic importance. In choice tests, adults preferred L. sinense for feeding and oviposition. These results show that A. tsekooni is relatively host specific and warrants further testing as a biocontrol agent of Chinese privet in the United States.     

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.     

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.     

广聚萤叶甲成虫取食经历对后续 产卵寄主选择的影响

【背景】取食经历对植食性昆虫的寄主选择行为具有较大影响,影响天敌昆虫寄主专一性测定的设计和结果解释。【方法】采用选择性试验,观察了入侵豚草的重要天敌——广聚萤叶甲成虫羽化后取食不同植物对其后续产卵寄主选择的影响。【结果】与取食豚草的试虫相比,有取食三裂叶豚草、苍耳或菊芋经历的成虫选择苍耳产卵的频次增加,不再对豚草表现出明显的选择偏好性。对产卵识别期的Cox模型分析结果表明,成虫早期取食不同植物,对后续产卵选择有显著影响,成虫羽化后如果先取食豚草或三裂叶豚草,则选择苍耳产卵的倾向显著低于豚草;但如果先取食苍耳、菊芋和农家向日葵,则选择苍耳产卵的倾向与豚草无显著差异。【结论与意义】由此推测,广聚萤叶甲初羽化成虫取食的植物对其后续产卵选择具有较大影响,因而在寄主专一性测定中应关注测试前饲喂的植物种类。     

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.