Effect of temperature on the development and consumption of Phaenochilus kashaya (Coleoptera: Coccinellidae), a predator of the cycad aulacaspis scale,Aulacaspis yasumatsui

The cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), is a serious pest of ornamental cycads in the southeastern United States. In Florida, CAS was first reported in Miami in 1996 and is now present in 43 Florida counties. Chemical control of CAS can be effective, but it is expensive, and insecticides must be frequently and regularly applied. Recent surveys of natural enemies in Thailand identified a new potential biological control agent of CAS. The objective of this study was to examine the temperature requirements and consumption rates of Phaenochilus kashaya Giorgi and Vandenberg (Coleoptera: Coccinellidae). Survival, developmental time and number of scales consumed by P. kashaya were measured at four constant temperatures (20, 25, 30 and 35°C). In addition, adult longevity and consumption were recorded at three constant temperatures (20, 25 and 30°C). Phaenochilus kashaya completed development to adulthood at temperatures ranging from 20 to 30°C. The highest survival rate (48%) was obtained at 25°C, and the accumulated degree-days for this species was 714. Phaenochilus kashaya is a voracious predator of CAS, consuming 380 scales during the larval stage and an estimated 4700 scales during the adult stage. Adult longevity varied from 59 days at 30°C to 220 days at 20°C for females, and 31 days at 30°C to 148 days at 20°C for males. The intrinsic rate of increase (r _m) was 0.34 at 20°C, which is high compared with other coccinellid species. According to this study, P. kashaya has the potential to be an effective biological control agent of the CAS in Florida.     

Life table of cycad scale,Aulacaspis yasumatsui (Hemiptera: Diaspididae), reared on Cycas in Taiwan

Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), an invasive scale insect, attacks different species of Cycas (Cycadales: Cycadaceae) in Taiwan. Development, survival and fecundity of A. yasumatsui were studied on Cycas taitungensis at 24 °C, 70 ± 10% RH, and a photoperiod of 12:12 (L:D) h under laboratory conditions. Data on the life history of A. yasumatsui were analyzed using the age-stage, two-sex life table, to address variable development rates among individuals and between sexes. The egg incubation time was 7.26 days for both females and males and female nymphal development duration was 28.65 days. The development duration of male nymphal stages + pupal stage was 19 days. The total pre-oviposition period (TPOP) was 35.92 days. The maximum longevity of female adults was 67 days and 1 day for males. The intrinsic rate of increase (r) was 0.100 day^-1, the finite rate of increase (λ) was 1.11 day^-1, the net reproduction rate (R_o) was 111.51 offspring /individual, and the mean generation time (T) was 47.24 day. Life table data can be used to project population growth, to design mass rearing programs and to establish management tactics to control insect pests.     

Two-sex life table and predation rate of Cybocephalus flavocapitis Smith (Coleoptera: Cybocephalidae) reared on Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), in Taiwan

Development, survival, and fecundity of the predator, Cybocephalus flavocapitis Smith, reared on its host Aulacaspis yasumatsui Takagi, were studied at 24 °C, 70 ± 10% RH, and photoperiod of 12:12 (L:D) h under laboratory conditions. Developmental periods for egg, larval, and pupal stages were 6.4, 12.1 and 14.3 d, respectively. The mean longevity of adults was 85.1 d. Raw life history data of all individuals were pooled and analyzed according to the age-stage, two-sex life table theory. The intrinsic rate of increase (r) was 0.05 d^? 1, the finite rate of increase (λ) was 1.05 d^? 1, the net reproductive rate (R₀) was 29.1 offspring, and the mean generation time (T) was 63.3 d. When we incorporated the age-stage variability of predation rate with the two-sex life table, our results indicated that the predation rates of larvae and adults were 743 eggs and 1595 scales, respectively. The net predation rate of C. flavocapitis is 1874 scales. Additionally, we compared life table results of C. flavocapitis with those of a previous life table study on Cybocephalus nipponicus because both species are currently used for the biological control of A. yasumatsui in Taiwan.     

Biology and preliminary host range assessment of two potential kudzu biological control agents

Two insect species from China, Gonioctena tredecimmaculata (Jacoby) (Coleoptera: Chrysomelidae) and Ornatalcides (Mesalcidodes) trifidus (Pascoe) (Coleoptera: Curculionidae), were studied in quarantine in the United States as potential biological control agents for kudzu, Pueraria montana variety lobata (Willd.) Maesen and S. Almeida. Adults of G. tredecimmaculata were ovoviviparous and reproduced throughout the summer, producing offspring that had an obligate adult diapause. In no-choice tests, adult and larval G. tredecimmaculata rejected most of the plant species tested, but consumed foliage and completed their life cycle on soybean (Glycine max L. Merr.) and on a native woodland plant, hog-peanut (Amphicarpaea bracteata L. Fernald), which are in the same subtribe as kudzu (Glycininae). Insects showed similar responses to field- and greenhouse-grown soybean and kudzu foliage, despite measurable differences in leaf traits: field-grown foliage of both plants had greater leaf toughness, higher total carbon content, higher trichome density, and lower water content than greenhouse foliage. O. trifidus adults also rejected most of the plants tested but fed on and severely damaged potted soybean and hog-peanut plants in addition to kudzu. Further tests in China are needed to determine whether these species will accept nontarget host plants under open-field conditions.     

茶椰圆蚧的重要天敌——日本方头甲及其捕食作用的研究

本研究了日本方头甲的种群动态,捕食习性、功能反应、干扰反应与农药毒性,结果表明,日本方头甲在福安社口茶园一年有４个发生高峰,与茶椰圆蚧二龄若虫的发生期跟随现象明显,对第二代茶椰圆蚧控制作用强。成虫和幼虫均或捕食各个虫态的茶椰圆蚧,成虫捕食具有日节律;雌成虫的捕食最大于雄成虫,雌、雄成虫捕食茶椰圆蚧的功能反应均为ＨｏｌｌｉｎｇⅡ型反应。雌成虫在１０℃下捕食率最低,１５℃次之,３５℃最高,在１０－３     

Evaluation of Alternaria crassa for biological control of Jimsonweed: Host range and virulence

An isolate of Alternaria crassa (Saac.) Rands was obtained from a naturally occurring leaf-spotting disease of Jimsonweed (Datura stramonium L.) plants. The fungus was induced to sporulate and used to reproduce the disease on seedlings in greenhouse and field inoculation tests. The fungus grew best at 25–35°C, and germinated best at 25–30°C. Infectivity on Jimsonweed was reduced at temperatures below 20°C and above 30°C. In greenhouse host range studies, conidia of the fungus were highly pathogenic to Jimsonweed, moderately pathogenic to ‘Marion’ and ‘Beefsteak’ tomatoes, and non-pathogenic to several other crop and weed species representing seven botanical families. These findings indicate that A. crassa has potential as a biological herbicide for controlling Jimsonweed.     

Host range of Secusio extensa (Lepidoptera: Arctiidae), and potential for biological control of Senecio madagascariensis (Asteraceae)

Secusio extensa (Lepidoptera: Arctiidae) was evaluated as a potential biological control agent for Madagascar fireweed, Senecio madagascariensis (Asteraceae), which has invaded over 400 000 acres of rangeland in the Hawaiian Islands and is toxic to cattle and horses. The moth was introduced from southeastern Madagascar into containment facilities in Hawaii, and host specificity tests were conducted on 71 endemic and naturalized species (52 genera) in 12 tribes of Asteraceae and 17 species of non‐Asteraceae including six native shrubs and trees considered key components of Hawaiian ecosystems. No‐choice feeding tests indicated that plant species of the tribe Senecioneae were suitable hosts with first instars completing development to adult stage on S. madagascariensis (78.3%), Delairea odorata (66.1%), Senecio vulgaris (57.1%), Crassocephalum crepidioides (41.2%), and at significantly lower rates on Emilia fosbergii (1.8%) and Erechtites hieracifolia (1.3%). A low rate of complete larval development also was observed on sunflower, Helianthus annuus (11.6%), in the tribe Heliantheae. However, sunflower was rejected as a potential host in larval‐feeding and adult oviposition choice tests involving the primary host S. madagascariensis as control. Although larvae died as first instars on most test species, incomplete development and low levels of feeding were observed on nine species in the tribes Heliantheae, Cardueae and Lactuceae. Larvae did not feed on any non‐Asteraceae tested, including species with similar pyrrolizidene alkaloid chemistry, crops, and six ecologically prominent native species. Because all species of Senecioneae are non‐native and weedy in Hawaii, these results indicate that S. extensa is sufficiently host‐specific for introduction for biological control. High levels of feeding damage observed on potted plants indicate that S. extensa can severely impact the target fireweed as well as D. odorata, a noxious weed in native Hawaiian forests.     

Host specificity of the two lace bugs,Dictyla echii andDictyla nassata [Hem.: Tingidae] potential biological control agents for paterson's curse (Echium plantagineum)

Two species of lace bugs,Dictyla echii (Schrank) andDictyla nassata (Puton), were tested for host specificity as potential biological control agents for Paterson's curse,Echium plantagineum L. A range of test plants from the family Boraginacea, from related families in the order Laminales and from species of economic importance in Australia which have had inadequate exposure in their countries of origin, were tested under both choice and non-choice conditions. These bugs proved to be oligophagous within the Boraginaceae and attacked a number of Australian native species. Because of this,D. echii andD. nassata are not considered sufficiently specific to release from quarantine and tests were abandoned before completion of the test plant list.

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Résumé La spécificité parasitaire deDictyla echii (Schrank) etD. nassata (Puton) a été étudiée en tant qu'agents de lutte biologique potentiels contreEchium plantagineum L. Dans les conditions de choix et d'absence de choix, une série de plantes ont été testées provenant de la famille des Boraginacées, des familles voisines de l'ordre des Laminales et d'espèces d'importance économique en Australie qui n'avaient pas fait l'objet de tests suffisants dans leur pays d'origine. Ces punaises se révèlent oligophages en ce qui concerne les Boraginacées et s'attaquent à de nombreuses espèces indigènes d'Australie. Pour cette raison,D. echii etD. nassata ne sont pas considérées comme suffisamment spécifiques pour les relacher à partir de la quarantaine et les tests ont été abandonnés avant d'épuiser la totalité des plantes prévues sur la liste.

     

Phytoseiid predators as potential biological control agents for Bemisia tabaci

Mites of the family Phytoseiidae are known to be predators of whiteflies in several agroecosystems, especially of Bemisia tabaci Gennadius, a pest with high resistance to chemical insecticides that occurs in greenhouses in temperate regions. We collected predatory mites that were found to co-occur with whiteflies in the Middle East for control of B. tabaci: Typhlodromus athiasae (Porath and Swirski), Neoseiulus barkeri Hughes, Typhlodromips swirskii (Athias-Henriot), Euseius scutalis (Athias-Henriot), Phytoseius finitimus Ribaga. As a first step in the evaluation of these species as biological control agents, we measured their life-history traits when feeding on whiteflies. The intrinsic rates of increase (r _m) of the predatory mite species ranged between 0.131 and 0.215 per day and E. scutalis had the highest r _m estimated. Comparisons with the r _m of B. tabaci indicate that some of the species should be capable of suppressing local populations of whitefly. The ability of predators to use alternative food was also tested, since it facilitates the setup of mass cultures and it can promote their persistence in the crop, even if the prey is scarce. All predatory mite species tested were able to survive and reproduce on a diet of broad bean pollen.     

Epiblema tetragonana and Epinotia ustulana (Lepidoptera: Tortricidae), two potential biological control agents for the invasive plant,Rubus ellipticus

Himalayan yellow raspberry, Rubus ellipticus is one of the world’s 100 worst invasive alien species. The plant has become a serious problematic weed in Hawaii, USA and is naturalized in many other countries. Screening of potential biological control agents is being conducted in its native region in Asia. In this paper, we report on the field distribution, abundance and host specificity of two leaf-rolling moth species, Epinotia ustulana and Epiblema tetragonana (Lepidoptera: Tortricidae). In larval non-choice tests both species only developed on plants in the genus Rubus. However, in adult choice oviposition tests, both leaf rollers showed a strong preference for R. ellipticus over other species. Furthermore, leaf-rolls by these two insects were only found on R. ellipticus in the field. These results indicate the moths have a narrow host range. Our field surveys also showed that both moth species are widely distributed in Yunnan Province, southeastern China, with up to 115 leaf rolls per plant, suggesting high levels of damage. These findings indicate that the two insects have considerable potential for biological control of R. ellipticus, though further host range tests should be conducted using more native plant species in Hawaii.     

Prioritizing areas in the native range of hygrophila for surveys to collect biological control agents

Native to Southeast Asia, Hygrophila polysperma is an invasive aquatic weed of lotic habitats in the southern United States and Mexico. An increase in the number of water bodies invaded by hygrophila since 1990 suggests that current methods employed to control this weed are inadequate. Classical biological control may be a viable option for long term regulation of hygrophila in the invaded range. In this study, we used the Maximum Entropy Species Distribution Model (MaxEnt) to prioritize climatically suitable native habitats in India and Bangladesh for conducting exploratory surveys to collect biological control agents. In total, 164 point occurrences from the United States and Mexico and 20 predictor variables, including 19 bioclimatic variables and altitude, were used to predict the native distribution of hygrophila. Performance of the model was statistically verified using threshold dependent binomial tests and area under the curve (AUC) score of the receiver operating characteristic (ROC) curve plot. The results showed that the model performed significantly better than random in both binomial tests and AUC analyses. High suitability of occurrence of hygrophila was predicted in the northeastern region of India and northern and eastern parts of Bangladesh. Based on percent omission of known native occurrences, a color-coded final distribution map was prepared to prioritize areas for conducting future surveys. Our study proposes a technique that can be useful for prioritizing areas in native ranges for exploratory surveys to collect biological control agents.     

Host specificity of the rust Phragmidium violaceurn, a potential biological control agent of European blackberry

The host specificity of the rust fungus Phragmidium violaceum, a potential biological control agent of European blackberry (Rubus fruticosus) was studied by inoculating a mixture of 15 isolates of the rust on 108 plants of importance to the Australasian region. A scale of infection types was developed based on the results of microscopic and macroscopic observations of the reaction of host and non-host plants to the rust. The results showed that P. violaceum has a limited host range in the genus Rubus. The rust was able to reproduce on 17 taxa of Rubus previously unrecorded as hosts, including Australasian species of Rubus subgenera Dalibarda and Lampobatus. All other taxa attacked were species of Rubus subgenus Eubatus and the majority were hybrid cultivars containing European blackberry species.     

Intraguild interactions between two biological control agents in citrus fruit: implications for biological control of medfly

The parasitoid wasp Spalangia cameroni and the predatory beetle Pseudoophonus rufipes have long been studied for use as biological control agents against the Mediterranean fruit fly Ceratitis capitata, particularly in citrus fruit orchards. Nevertheless, these two species of natural enemies, when competing for a common resource, may experience intraguild predation (IGP) interactions. These possible interactions, affecting parasitism and predation, have been evaluated in the present work, under laboratory conditions, through potential changes in functional response. Regarding host/prey density, both natural enemies, when acting alone, showed a type II functional response. Nevertheless, due to IGP, S. cameroni, in the presence of P. rufipes, showed a higher fertility rate and a type III functional response. The parasitism behaviour of S. cameroni was affected by the presence of the predator, reducing the host handling time. Conversely, the parasitism rate of S. cameroni did not vary in the presence of P. rufipes but the degree of superparasitism decreased and led to an increased fertility rate and an increasingly female‐biased sex ratio. Meanwhile, the predatory efficiency of P. rufipes was not affected by the presence of S. cameroni but discrimination between parasitised and unparasitised pupae of C. capitata, with a preference for the latter, was reported for this predator species. Our results suggest that in biological control programmes, the use of only one of these species is recommended at low infestation levels, whilst at high densities of the pest, the combination of both natural enemies seems to be the most appropriate strategy.     

Biology and field host range of Ceutorhynchus cardariae,a potential biological control agent for Lepidium draba

Lepidium draba (Brassicaceae) is a clonal herb, originating from Eurasia, which is invasive in North America. A classical biological control project was initiated in 2001, and the gall‐forming weevil Ceutorhynchus cardariae was prioritized as a candidate agent. We studied its biology and field host range between 2003 and 2014 in the laboratory and a common garden in Switzerland and in the field in Romania. Ceutorhynchus cardariae is a univoltine to bivoltine species. In Switzerland, oviposition usually started at the beginning of March and can occur at temperatures as low as 2.5°C. Galls are formed on stems, leaf stalks and midribs of L. draba rosettes and bolting plants. Gall size increased with an increasing number of larvae per gall. The three larval instars feed inside the galls and leave the plant to pupate in the soil once mature. In Switzerland, development from egg to adult took about 12 weeks in spring. Adults emerged from May to July. After a brief feeding period, adults aestivate. From late summer, feeding recommenced and females may oviposit, forming a partial second generation. Eggs and all larval instars can be found in galls throughout winter. The rate of larval ectoparasitism reached 78%, while endoparasitism was low with a maximum of 2.3%. Lepidium draba populations differed in their suitability for development (number of C. cardariae produced), indicating that effectiveness of C. cardariae – in case released – may be variable. In the field, we observed that gall formation by C. cardariae can severely stunt or even kill shoots. Investigations on the field host range of C. cardariae indicated that only the closely related Lepidium campestre may act as an alternative host for the weevil in Europe. Host‐specificity tests are underway to determine its environmental safety before field release in North America is being considered.     

A testing sequence for reducing rejection of potential biological control agents for weeds

Many insect agents, selected for the biological control of weeds and tested under restricted cage conditions, have been rejected as unsafe for introduction because of an apparent increase in host range compared with that observed under natural conditions. These aberrant results appear to be due to the insects' inability to follow the normal behavioural sequence based on appropriate cues which lead to correct host selection. It is suggested here that a reverse sequence of testing which progressively reduces the degree of restriction and deletes unattacked plants at each stage, until only a few remain to be tested under conditions as near natural as possible, would allow previously rejected insects to be used as biological control agents.     

蚧虫的病原真菌及其在生物防治中的潜力

蚧虫（半翅目：蚧总科）是农林果树和花卉的一类重要害虫。作者综述了寄生蚧虫的虫生真菌及其在生物防治中的潜力。总结了昆虫病原真菌作为生物杀虫剂的研究历史,并将其划分为3个发展阶段,即开创阶段、缓慢发展阶段和快速发展阶段。讨论了该领域在中国的研究现状。列出了世界上目前已记录的蚧虫病原真菌,包括55属140种,及其寄主蚧虫的名录。对虫生真菌未来的研究和开发提出了4点建议。     

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 range testing of a prospective classical biological control agent against cabbage maggot,Delia radicum,in Canada

The introduction of a European natural enemy, Aleochara bipustulata L. (Coleoptera: Staphylinidae), is being considered for control of cabbage maggot, Delia radicum (L.) (Diptera: Anthomyiidae) in canola in Canada, and the host specificity of this pupal parasitoid must first be studied. Contemporary guidelines were used to select 18 species of Diptera to represent non-target species taxonomically related or ecologically similar to reported hosts of A. bipustulata, or beneficial species. No-choice tests were used to determine which of the 18 species are within A. bipustulata’s fundamental host range, and whether puparial structure or mass or duration of pupal development influence their acceptability and suitability as hosts. Five species were consistently suitable as hosts, and these were either relatively small or were taxonomically closely related to the target host. The probability that a puparium would be entered by a parasitoid larva was greatest for small puparia, but was unaffected by pupal duration. The probability of completing parasitoid development once a puparium was entered was influenced by both puparial mass and duration of pupal development. Pitfall traps to assess habitat associations caught A. bipustulata adults in a variety of crop habitats but none were caught in forests. Host range and habitat data are used to argue that there is little risk of parasitism to beneficial taxa. Non-target species taxonomically closely related to the target D. radicum or with small puparia may fall within the fundamental host range of A. bipustulata. However, risk to many of these species may be minimal because of the habitat preferences of the parasitoid and its cues for host-finding and recognition.     

Evaluation of biogeographical factors in the native range to improve the success of biological control agents in the introduced range

Biogeographical factors associated with the invasive weed Arundo donax were evaluated at 22 different locations in four countries in its native range in reference to its key herbivore, an armoured scale insect, Rhizaspidiotus donacis. Data on climate, plant health and quality, soil conditions and anthropogenic influences were analysed for each site and compared to populations of R. donacis. Climate modelling from location data in Spain and France accurately predicted the native range of the scale in the warmer, drier parts of Italy and Greece and was used to predict its distribution in North America. Presence of the scale was not associated with specific soil type or characteristics. However, it was positively associated with a higher percentage of dead stems and significantly lower biomass of A. donax. Micronutrient sampling using leaf material found that sulphur was negatively correlated with aluminium and positively correlated with boron for scale density. Disturbance of field sites by cutting and/or addition of supplemental irrigation during summer appeared to disrupt the synchronised seasonal phenology of A. donax and R. donacis, leading to more robust stands. These biogeographical factors from the native range indicate that R. donacis should have the greatest impact in warm, dry climates in the introduced range where A. donax is undisturbed.