Temperature-dependent development and potential distribution of Episimus utilis (Lepidoptera: Tortricidae), a candidate biological control agent of Brazilian peppertree (Sapindales: Anacardiaceae) in Florida

The invasive Brazilian peppertree (Schinus terebinthifolius Raddi), native to South America, is widely established throughout central and south Florida. The defoliating leaflet-roller Episimus utilis Zimmerman was selected as potential biocontrol agent of this invasive species. The objectives of this study were to determine development rate and survival of E. utilis at seven constant temperatures (10, 15, 20, 25, 30, 33, and 35 degrees C) and generate prediction maps of the number of generations per year this species may exhibit in the United States. The rate of development of E. utilis as a function of temperature was modeled using linear regression to estimate a lower developmental threshold of 9.6 degrees C and the degree-day requirement of 588. The Logan nonlinear regression model was used to estimate an upper developmental threshold of 33 degrees C. Cold tolerance of E. utilis was examined using all insect stages, and each stage was exposed to three constant temperatures (10, 5, 0 degrees C) for 0.5, 1, 2, 4, and 8 d (or until all insects died). The pupal stage was the most cold tolerant with 100% mortality after 12 d at 0 degrees C. The pupal lethal times at 5 (Ltime50 = 10 d, Ltime90 = 28 d) and 0 degrees C (Ltime50 = 5 d, Ltime90 = 9 d) were used to generate isothermal lines to predict favorable regions for E. utilis establishment. A GIS map was generated to predict the number of generations of E. utilis (range, 0.5-9.8) across all Brazilian peppertree range in the United States. The potential for establishment of E. utilis and its probable distribution in the continental United States was examined.     

Biology and host range of Omolabus piceus, a weevil rejected for biological control for Schinus terebinthifolius in the USA

Surveys for biological control agents of the invasive weed Schinus terebinthifolius (Anacardiaceae) discovered two Omolabus weevils (Coleoptera: Attelabidae) feeding on the plant in its native range. Molecular and morphological analysis indicated that one of these species consistently fed on the target weed and the other species fed more broadly. Aspects of the biology and host range of the more specific species, Omolabus piceus (Germar) were examined to determine its suitability as a biological control agent of S. terebinthifolius in the USA. Adults feed on newly formed leaves, and eggs, larvae and pupae develop in curled fragments of leaves, called nidi. Larvae consumed an average of 11.3 (±0.4) mg throughout their development which required 15.1 (±0.2) days. An average of 31.6 (±2.7) eggs were laid per female during their 23.8 (±2.2) day lifetime, after a 3.4 (±1.0) day preoviposition period. In no-choice tests, O. piceus adults fed and oviposited on all tested native North American, Caribbean and agricultural Anacardiaceae species except for M. indica. The field host-range of O. piceus, as determined by samples of host use in the native range, included three Schinus, two Lithrea and one Anacardium species. Therefore, we do not recommend O. piceus for biological control of S. terebinthifolius in the USA. However, the utilization of this species in other infested areas such as Hawaií and Australia should be considered.     

Biology, host specificity tests, and risk assessment of the sawfly Heteroperreyia hubrichi, a potential biological control agent of Schinus terebinthifolius in Hawaii

Heteroperreyia hubrichiMalaise (Hymenoptera: Pergidae), a foliagefeeding sawfly of Schinusterebinthifolius Raddi (Sapindales:Anacardiaceae), was studied to assess itssuitability as a classical biological controlagent of this invasive weed in Hawaii. No-choice host-specificity tests were conductedin Hawaiian quarantine on 20 plant species in10 families. Besides the target weed, adultfemales oviposited on four test species. Females accepted the Hawaiian native Rhussandwicensis A. Gray (Sapindales:Anacardiaceae) as an oviposition host equallyas well as the target species. The other threespecies received significantly fewer eggs. Neonate larvae transferred onto test plantssuccessfully developed to pupae on S.terebinthifolius (70% survival) and R.sandwicensis (1% survival). All other 18test plant species failed to support larvaldevelopment. A risk analysis was conducted toquantify the acceptability of non-targetspecies as host plants for H. hubrichi onthe basis of the insect's performance atvarious stages in its life cycle. Risk ofdamage to all plant species tested wasinsignificant except for R. sandwicensis. Risk to this native plant relative to S.terebinthifolius was estimated at 1%. Currently this level of risk is too high torequest introduction of this insect into theHawaiian environment. Detailed impact studiesin the native range of S. terebinthifoliusare needed to identify thepotential benefit that this insect offers. Also, field studies in South America withpotted R. sandwicensis would give a morereliable analysis of the risk this nativeHawaiian plant would face from naturalpopulations of H. hubrichi.     

Biology and host range of Platyphora semiviridis, a leaf beetle evaluated as a potential biological control agent for Solanum mauritianum in South Africa

The South American tree Solanum mauritianum is a major environmental weed in the high-rainfall regions of South Africa and has been targeted for biological control. Potential agents included five species of the genus Platyphora, which were imported from South America in 1994. Platyphora species associated with Solanaceae reputedly have very specific habitat requirements and host plant preferences in the field. Despite this, host-specificity tests on one species, Platyphora semiviridis, revealed a broad physiological host range. Although laboratory tests showed that P. semiviridis is confined to Solanum species and cannot survive on solanaceous crops outside that genus, it developed on potato and cultivated eggplant (aubergine) as well as on 10 native South African Solanum species. With few exceptions, there were no consistent differences in survival and duration of development on these compared with S. mauritianum. Furthermore, at least six of these non-target species, including potato and eggplant, supported breeding colonies of the beetles in cages. During choice tests in both small and larger cages, P. semiviridis avoided potato but did not consistently discriminate between S. mauritianum, eggplant and six native solanums for larviposition. Despite these findings, P. semiviridis has never been recorded on either potato or eggplant in South America, where it was only observed to feed on S. mauritianum. Although there are several reasons why P. semiviridis is unlikely to attack non-target Solanum species in the field, it will not be released in South Africa because there are other imported agents which have displayed narrower physiological host ranges and which may be more effective.     

Biology and host range of Heterapoderopsis bicallosicollis: a potential biological control agent for Chinese tallow Triadica sebifera

Chinese tallow, Triadica sebifera, is an invasive weed that infests natural and agricultural areas of the south-eastern USA. A candidate insect for biological control of Chinese tallow has been studied under quarantine conditions. The biology and host range of a primitive leaf-feeding beetle, Heterapoderopsis bicallosicollis, were examined between July 2008 and February 2010. H. bicallosicollis adults fed on 16 of 29 non-target species and survived >10 days on all 29 species, more than the number of days that adults survived when deprived of food. Additionally, adult feeding, oviposition and partial larval development occurred on a Florida endangered species, Heterosavia bahamensis. These data indicated that H. bicallosicollis adult feeding and oviposition may not be limited to the target weed and included several valued North American natives. Therefore, as its release could pose unacceptable risks, testing of this species was discontinued and the quarantine colony destroyed.     

Pyemotes tritici (Acari: Pyemotidae), a potential biological control agent of Anagasta kuehniella (Lepidoptera: Pyralidae)

The feasibility of utilizing Pyemotes tritici as a biological control agent against the Mediterranean flour moth, Anagasta kuehniella was investigated. The results from experiments designed to assess the effect of P. tritici on cohorts of A. kuehniella of different ages were not significant. They suggested that moth larvae were most severely affected if physogastric P. tritici females were introduced 10 days after moth eggs were added to the experimental arena. In part II, the impact of various densities of P. tritici on flour moths indicated that complete destruction of cohorts of 400 A. kuehniella larvae was achieved by adding about 80 female P. tritici 10 days after the eggs were laid or 6 days after the larvae hatched. Half the above density of pyemotid mites gave inconsistent, but at times complete control.     

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.     

Biology and host range of the leafminer,Pseudonapomyza sp. (Diptera: Agromyzidae), a potential biological control agent for Tecoma stans (Bignoniaceae) in South Africa

Tecoma stans (Bignoniaceae), is an evergreen shrub that has a wide natural distribution in the tropical and subtropical parts of the western hemisphere. This shrub is native to Mexico and the southern regions of the USA. This weed is widely distributed in South Africa and neighbouring countries. As part of the biological control initiative, a leafmining fly, Pseudonapomyza sp. (Diptera: Agromyzidae), was imported into South Africa, and was subsequently studied as a potential biological control agent for T. stans. During no-choice tests involving 46 plant species in 16 families, Pseudonapomyza sp. only oviposited and developed on T. stans. Neither oviposition nor larval development was recorded on the closely related and indigenous plant species. When six plant species in the Bignoniaceae family were exposed to Pseudonapomyza sp. during multi-choice tests, oviposition and larval development only occurred on T. stans. It was concluded that Pseudonapomyza sp. was sufficiently host-specific to be released against T. stans in South Africa. Pseudonapomyza sp. also displayed very promising biological attributes that could enhance its effectiveness to control T. stans.     

Biology of Mastrus ridibundus (Gravenhorst), a potential biological control agent for area‐wide management of Cydia pomonella (Linneaus) (Lepidoptera: Tortricidae)

The codling moth Cydia pomonella (Linnaeus) (Lepidoptera: Tortricidae) is a serious pest of pome fruit crops. A natural enemy of codling moth, the larval ectoparasitoid Mastrus ridibundus (Gravenhorst) (Hymenoptera: Ichneumonidae) has been imported into South America from the USA but little is known about the biology and ecology of the wasp, knowledge that is needed to design an efficient strategy of release and establishment. Experiments were carried out to assess important traits of the biology of the parasitoid in relation to its possible use as a biocontrol agent for codling moth. When M. ridibundus females were offered larvae ranging in weight from 37 to 78 mg, they oviposited more eggs on heavier hosts. In another study, the adult wasps were offered honey, diluted honey (10%) or pollen in paired choice tests and both males and females preferred honey over the other two foods. Females preferred 10% honey over pollen, while the males showed the opposite preference. Honey‐fed females lived longer than starved females. Adults died rapidly at 35°C, while they lived 20 days at 25°C and 12–17 days at 15°C. Female wasps had on average 25 ± 14 and 18 ± 11 progeny at 15 and 25°C, respectively, but they did not had progeny at 35°C. The development time (egg to adult emergence) was on average 44 ± 7 and 24 ± 2 days at 15 and 25°C respectively. Immature insects did not reach the adult stage at 35°C.     

Biology ofAllothrombium pulvinum (Acariformes: Trombidiidae), a potential biological control agent of aphids in China

Allothrombium pulvinum Ewing is a common natural enemy of aphids and other small arthropods in China. It is univoltine species. The eggs hatch in spring, nymphs emerge in early summer, and adults emerge in autumn. Larvae are ectoparasites of aphids, whereas deutonymphs and adults are free-living predators of aphids, spider mites and some lepidopterous eggs. Adults hibernate in soil during winter and females lay their eggs in spring. In the laboratory the duration of immature stages is 74 days at 20–30°C. Larvae kill their host within 1–3 days when the mite load of the host is 2 or more. They decrease the reproductive rate of adult aphids and arrest the development of early-instar aphid nymphs when the mite load is 1. Larvae are capable of controlling the host population growth when the level of parasitism is high. Owing to its wide occurrence,A. pulvinum could be a potential candidate in the biological control of aphids. It is suggested that its impact on the host pest be evaluated in IPM programs in the future.     

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.     

Life cycle and host range of the gall-forming moth,Meskea horor (Lep.: Thyrididae), and its suitability as a biological control agent forSida acuta andS. rhombifolia (Malvaceae)

The life cycle of the moth,Meskea horor Dyar and its host specificity toSida acuta andS. rhombifolia were investigated. Moths emerge early in the morning and are usually inactive during the day. Feeding by the adults in the field was not observed. However, those in the laboratory fed on flowers ofS. acuta, S. rhombifolia andHibiscus sabdariffa, and drank water or dilute honey solution. Eggs are mainly laid on the underside of leaves. Larvae ofM. horor form galls in the stems ofS. acuta andS. rhombifolia which retard plant growth and flowering. Pupation occurs within the gall. The development time from egg to adult was 192 days and the adults lived for 11 to 14 days. Forty-eight plant species were tested to determine the host range ofM. horor. Adults emerged from seven species of plants in the family Malvaceae and larvae formed galls but died before pupating on a further 17 species.M. horor is considered to have too broad a host range to be used as a biological control agent forS. acuta andS. rhombifolia.     

Nuclear polyhedrosis virus as a biological control agent for Malacosoma americanum (Lepidoptera: Lasiocampidae)

In addition to damaging trees, the eastern tent caterpillar is implicated in early fetal loss and late‐term abortion in horses. In a field study, we evaluated the potential biological control of the caterpillar using eastern tent caterpillar nuclear polyhedrosis virus (ETNPV), a naturally occurring virus that is nearly species‐specific. Egg masses were hatched and second instar larvae were fed virus‐inoculated foliage to propagate the virus in vivo. Then, a viral pesticide was formulated at concentrations of 10⁴, 10⁶ and 10⁸ polyhedral inclusion bodies per ml. The pesticide was applied to foliage on which second, third and fourth instar caterpillars were feeding. When the majority of surviving larvae reached the sixth instar, colonies were collected and the surviving caterpillars counted. Mean numbers of surviving caterpillars per treatment were compared via 95% bootstrap confidence intervals. The data indicate second instar caterpillars were highly susceptible to the virus, but only at the highest concentration tested. Third instar caterpillars were also somewhat susceptible to high virus concentrations, while fourth instar caterpillars were fairly resistant. Our data provide the strongest evidence to date that ETNPV can be propagated, harvested and refined for formulation as a biological control agent for eastern tent caterpillar. Its use on this insect may be merited in circumstances where landowners and managers need to protect trees and horses.     

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16′ S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.     

Biology and host range of Ophiomyia camarae, a biological control agent for Lantana camara in South Africa

The herringbone leaf-mining fly, Ophiomyiacamarae Spencer, is a promising candidateagent for the biological control of the alieninvasive weed Lantana camara L. in SouthAfrica. During extensive host-specificity testsinvolving 39 test plant species from 12families, survival to adulthood was restrictedto L. camara, L. trifolia, and fourspecies of the closely-related genus Lippia (Verbenaceae) in no-choice tests. However, survival of the immature stages wassignificantly lower on L. trifolia andthe four Lippia species than on L.camara. In addition, O. camaraedisplayed very strong oviposition preferencefor L. camara during paired-choice tests.This narrow laboratory host range suggests thatO. camarae will not pose any risks tonon-target verbenaceous plants if released inSouth Africa. Permission to release O. camaraeinto South Africa was approved by the regulatoryauthorities in 2001.     

Introduction, establishment, and potential geographic range of Carmenta sp. nr ithacae (Lepidoptera: Sesiidae), a biological control agent for Parthenium hysterophorus (Asteraceae) in Australia

Parthenium (Parthenium hysterophorus L.), a major weed causing economic, environmental, and human and animal health problems in Australia and several countries in Asia, Africa, and the Pacific, has been a target for biological control in Australia since the mid-1970s. Nine species of insects and two rust fungi have been introduced as biological control agents into Australia. These include Carmenta sp. nr ithacae, a root feeding agent from Mexico. The larvae of C. sp. nr ithacae bore through the stem-base into the root where they feed on the cortical tissue of the taproot. During 1998-2002, 2,816 larval-infested plants and 387 adults were released at 31 sites across Queensland, Australia. Evidence of field establishment was first observed in two of the release sites in central Queensland in 2004. Annual surveys at these sites and nonrelease sites during 2006-2011 showed wide variations in the incidence and abundance of C. sp. nr ithacae between years and sites. Surveys at three of the nine release sites in northern Queensland and 16 of the 22 release sites in central Queensland confirmed the field establishment of C. sp. nr ithacae in four release sites and four nonrelease sites, all in central Queensland. No field establishment was evident in the inland region or in northern Queensland. A CLIMEX model based on the native range distribution of C. sp. nr ithacae predicts that areas east of the dividing range along the coast are more suitable for field establishment than inland areas. Future efforts to redistribute this agent should be restricted to areas identified as climatically favorable by the CLIMEX model.     

Papilio aegeus Donovan (Lepidoptera: Papilionidae) host plant range evaluated experimentally on ancient angiosperms

Abstract  Chemical similarities among ancient Angiosperms presumably played a role in the ecological and evolutionary diversification of the swallowtail butterflies (Papilionidae). The abilities of neonate larvae of the Citrus swallowtail, Papilio (= Princeps ) aegeus (from Queensland, Australia), to eat, survive and grow on leaves (a choice of young and old) of 34 plant species from families of ancient Angiosperms; 8 Rutaceae, 3 Magnoliaceae, 13 Lauraceae, 3 Monimiaceae, 1 Aristolochiaceae, 2 Apiaceae, 1 Sapotaceae, 1 Winteraceae and 2 Annonaceae were tested. It was apparent that there is genetic variation in populations of Rutaceae-specialised Australian P. aegeus for acceptance, consumption and larval growth, reflecting differential suitability of some native Australian Lauraceae species as food plants (as well as certain Winteraceae, Monimiaceae and non-Australian Magnoliaceae, Lauraceae and Annonaceae). No consumption or survival of P. aegeus was seen on Aristolochia elegans (Aristolochiaceae) or Pouteria australis (Sapotaceae) despite literature records alluding to this possibility. The Rutaceae specialist P. aegeus appears to have the fundamental detoxification capabilities for processing many existing species of the basal Angiosperm families, without having direct ancestors that historically had fed on them.     

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.     

Life history and host range of Alagoasa extrema (Coleoptera: Chrysomelidae: Alticinae), a potential biological control agent of Lantana spp. (Verbenaceae) in Australia

The life history and host range of the lantana beetle, Alagoasa extrema, a potential biocontrol agent for Lantana spp. were investigated in a quarantine unit at the Alan Fletcher Research Station, Brisbane, Australia. Adults feed on leaves and females lay batches of about 17 eggs on the soil surface around the stems of plants. The eggs take 16 days to hatch and newly emerged larvae move up the stem to feed on young leaves. Larvae feed for about 23 days and there are three instars. There is a prepupal non-feeding stage that lasts about 12 days and the pupal stage, which occurs in a cocoon in the soil, lasts 16 days. Teneral adults remain in the cocoon for 3 days to harden prior to emergence. Males live for about 151 days while females live for about 127 days. The pre-oviposition period is 19 days. In no-choice larval feeding trials, nine plant species, representing three families, supported development to adult. Three species, Aloysia triphylla, Citharexylum spinosum and Pandorea pandorana were able to support at least two successive generations. These results confirm those reported in South Africa and suggest that A. extrema is not sufficiently specific for release in Australia. Furthermore, it is not recommended for release in any other country which is considering biological control of lantana.