Defoliating broad-nosed weevil,Plectrophoroides lutra; not suitable for biological control of Brazilian pepper (Schinus terebinthifolius)

Adults of the weevil Plectrophoroides lutra were evaluated for biological control of Schinus terebinthifolius. No-choice tests conducted on North American and other valued plants indicated the adults fed on all species offered. Thus, it is unlikely that P. lutra will be developed as a biological control for S. terebinthifolius.     

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.     

Life history and host range of the leaf blotcher Eucosmophora schinusivora: a candidate for biological control of Schinus terebinthifolius in the USA

The host range of Eucosmophora schinusivora Davis and Wheeler (Lepidoptera: Gracillariidae) was studied to assess its suitability as a biological control agent of Schinus terebinthifolius Raddi (Anacardiaceae), a serious environmental and agricultural weed in the USA and elsewhere in the world. The life history of this insect species and its host range were determined in the laboratory with adult no-choice oviposition and larval development tests. This species has five instars, the first three are sap-feeding miners and the last two are tissue feeding. Total development time was 31.7 days. To examine specificity of this species, 10 plant species in Anacardiaceae were selected based on taxonomic relatedness to S. terebinthifolius, economic importance, and availability. In the laboratory, except for Anacardium occidentale and Cotinus obovatus, all of the tested species were accepted for oviposition with a marked preference for the weed S. terebinthifolius, Schinus molle, Rhus copallinum, Rhus sandwicensis and Pistacia chinensis. Complete development, from egg to adult, was achieved only on S. terebinthifolius, S. molle, R. copallinum, P. chinensis and Metopium toxiferum. In conclusion, E. schinusivora will not be considered for the biological control of S. terebinthifolius in the continental USA. However, the utilisation of this species in other infested areas such as Hawai'i and Australia should be considered.     

Comparison of two populations of Pseudophilothrips ichini (Thysanoptera: Phlaeothripidae) as candidates for biological control of the invasive weed Schinus terebinthifolia (Sapindales: Anacardiaceae)

Brazilian peppertree, Schinus terebinthifolia Raddi (Sapindales: Anacardiaceae) (hereafter Schinus), is one of the worst invasive species in Florida and Hawaii. The thrips Pseudophilothrips ichini Hood (Thysanoptera: Phlaeothripidae) is being considered as a potential biological control agent of Schinus. Two populations of this thrips were collected in the weed's native range; one from central-east Brazil (Ouro Preto thrips) and a second from north-east Brazil (Salvador thrips). Temperature requirements, adult fecundity and impact on different plant haplotypes by P. ichini were examined in the laboratory. Complete development of thrips from both populations occurred at temperatures ranging from 20 to 30°C. Two approaches were used to model the predicted distributions of the thrips populations in the USA: the physiological model (NAPPFAST) based on cold tolerance and the ecological niche model based on climatic variables (MaxEnt). The physiological model predicted that both populations of P. ichini may establish in similar areas of the USA, overlapping with the distribution of Schinus. However, the niche model predicted that only the Ouro Preto thrips could establish in the USA. The difference in model predictions suggests an apparent preadaptation of the Salvador thrips to lower temperatures than those experienced at the locations they were collected in Brazil. The Ouro Preto thrips had similar fecundity on two Florida Schinus haplotypes, whereas lower fecundity on haplotype A was found for the Salvador thrips. Based on these results, the Ouro Preto population may be better adapted to the climatic conditions and plant haplotypes found in Florida. Moreover, greenhouse studies indicated that Schinus growth was greatly reduced by thrips feeding, which may result in lower weed reproduction and densities in the field.     

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.     

Control of Pseudophilothrips ichini (Thysanoptera: Phlaeothripidae) with acephate to exclude a biological control agent of Schinus terebinthifolia

Post-release monitoring of biological control agents to determine impact on the target weed has recently received increased priority. Several methods are available to measure the impact of a biological control agent by manipulating the agent population while measuring fitness of the weed. Brazilian peppertree, Schinus terebinthifolia is one of the most damaging weeds in subtropical areas of Florida and Hawaii. A biological control agent, the thrips, Pseudophilothrips ichini is a sap-feeder that shows high levels of specificity and causes severe distortion of leaf tips of the weed. Thrips populations of this species and a generalist thrips, the red banded thrips Selenothrips rubrocinctus were experimentally manipulated by applications of the systemic insecticide acephate by both foliar applications and by an inserted encapsulated formulation. Foliar applications protected plants against red banded thrips for 29 days and against the biological control thrips, P. ichini for 22 days after treatment. Control with inserts were initially low but was achieved after 60 days and this control continued for 182 days after treatment. Manipulation of these biological control thrips populations with foliar or inserted formulations will assist in the determination of biological control agent impact.     

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 Tecmessa elegans (Lepidoptera: Notodontidae), a leaf-feeding moth evaluated as a potential biological control agent for Schinus terebinthifolius (Sapindales: Anacardiaceae) in the United States

During surveys for natural enemies that could be used as classical biological control agents of Schinus terebinthifolius Raddi (Brazilian pepper), the caterpillar, Tecmessa elegans Schaus (Lepidoptera: Notodontidae), was recorded feeding on the leaves of the shrub in South America. The biology and larval and adult host range of this species were examined to determine the insect's suitability for biological control of this invasive weed in North America and Hawaii. Biological observations indicate that the larvae have five instars. When disturbed, the late instar larvae emit formic acid from a prothoracic gland that may protect larvae from generalist predators. Larval host range tests conducted both in South and North America indicated that this species feeds and completes development primarily on members of the Anacardiaceae within the tribe Rhoeae. Oviposition tests indicated that when given a choice in large cages the adults will select the target weed over Pistacia spp. However, considering the many valued plant species in its host range, especially several North American natives, this species will not be considered further for biological control of S. terebinthifolius in North America.     

Biology and host range of the stem-boring beetle Aphanasium australe, a promising agent for the biological control of Hakea sericea in South Africa

The invasive Australian shrubHakea sericea Shrader is a majorenvironmental weed in the Western and EasternCape Provinces of South Africa. Dense,impenetrable thickets severely threaten theunique endemic vegetation of the Cape FloristicKingdom, increase fire hazards and reduce wateryields in catchments. Biological control,initiated in the 1970s, is largely confined tothe use of seed-feeding insect agents. Becausenone of these agents reduce the density ofexisting hakea populations, a stem-boringbeetle, Aphanasium australe (Boisduval) (Cerambycidae),was imported into quarantine in South Africa in1975. During multichoice oviposition tests,involving 12 Australian and six South Africanproteaceous species, in a walk-in cage, A. australe only oviposited on four species ofHakea and two exotic species of Grevillea. However, culturing difficultiesresulted in the suspension of host-specificitytesting after three years. Testing was resumedfollowing re-importations in 1995, also becauseA. australe also attacks Hakeagibbosa (Sm.) Cav., which is not attacked byany of the existing biocontrol agents. Duringno-choice survival tests, involving 66test plant species from 15 families, A. australe only developed on H. sericea, H. gibbosa and one exoticspecies of Grevillea. The contention thatA. australe is confined to the genus Hakea was confirmed by host records andsurveys in Australia which provided no evidenceof attacks on crop, pasture or related plants.The regulatory authorities accepted theseresults and A. australe was cleared forrelease in South Africa during 2001.     

Preliminary studies on Ramularia crupinae sp. nov. as a potential biological control agent for common crupina (Crupina vulgaris) in the USA

Common crupina, Crupina vulgaris (Asteraceae), is an alien weed of pastures and disturbed noncrop lands in the western USA. Research is underway in Europe to search for the natural enemies to control the weed. Among various natural enemies attacking C vulgaris, a new fungal species, Ramularia crupinae, has been found to cause serious damage to the weed. The fungus has been grown on various media and a modified V8 juice agar has proved to be suitable for both growth and sporulation. Greenhouse and field studies have shown that R. crupinae can kill infected leaves, thus reducing the plant growth, and in severe cases can kill the young plants. During preliminary host-range tests using various members of the Asteraceae, including artichoke and safflower, the fungus attacked only C. vulgaris.     

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.     

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.     

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.     

Winter climate change and the poleward range expansion of a tropical invasive tree (Brazilian pepper—Schinus terebinthifolius)

Winter climate change is expected to lead to the tropicalization of temperate ecosystems, where tropical species expand poleward in response to a decrease in the intensity and duration of winter temperature extremes (i.e., freeze events). In the southeastern United States, freezing temperatures control the northern range limits of many invasive nonnative species. Here, we examine the influence of freezing temperatures and winter climate change on the northern range limits of an invasive nonnative tree—Schinus terebinthifolius (Brazilian pepper). Since introduction in the 1800s, Brazilian pepper has invaded ecosystems throughout south and central Florida to become the state's most widespread nonnative plant species. Although Brazilian pepper is sensitive to freezing temperatures, temperature controls on its northern distribution have not been adequately quantified. We used temperature and plant occurrence data to quantify the sensitivity of Brazilian pepper to freezing temperatures. Then, we examined the potential for range expansion under three alternative future climate scenarios (+2°C, +4°C, and +6°C). Our analyses identify a strong nonlinear sigmoidal relationship between minimum temperature and Brazilian pepper presence, with a discrete threshold temperature occurring near ?11°C. Our future scenario analyses indicate that, in response to warming winter temperatures, Brazilian pepper is expected to expand northward and transform ecosystems in north Florida and across much of the Gulf of Mexico and south Atlantic coasts of the United States. These results underscore the importance of early detection and rapid response efforts to identify and manage the northward invasion of Brazilian pepper in response to climate change. Looking more broadly, our work highlights the need to anticipate and prepare for the tropicalization of temperate ecosystems by tropical invasive species.     

Biology and host specificity ofWeiseana barkeri (Col.: Chrysomelidae): A biological control agent forAcacia nilotica (mimosaceae)

The leaf-feeding beetleWeiseana barkeri Jacoby feeds onAcacia nilotica (L.) Willdenow ex Delile in Kenya. Host specificity tests show it is host specific toA. nilotica and approval has been given for its field release in northwest Queensland. A novel rearing and host specificity-testing technique is reported whereA. nilotica foliage stimulates oviposition into strips of corrugated cardboard.     

Failure of the introduction ofActinote anteas (Lep.: Acraeidae) from Costa Rica as a biological control candidate forChromolaena odorata (Asteraceae) in South Africa

Actinote anteas from Costa Rica was screened as a biological control candidate forChromolaena odorata in South Africa. Preliminary starvation trials suggest thatA. anteas is species specific. There are seven larval instars and the life cycle is completed in 101–169 days. The culture died out after three generations possibly because of incompatibility with the form of the local species ofC. odorata or disease.     

Host range testing and life history of the defoliator Hymenomima nr. memor: an unsuitable biological control agent for Schinus terebinthifolia in the U.S.A.

Host range of larvae of Hymenomima nr. memor (Lepidoptera: Geometridae) was examined in quarantine to evaluate its suitability as a biological control of Brazilian peppertree, Schinus terebinthifolia. Brazilian peppertree, S. terebinthifolia is an environmental and agricultural weed from South America that had invaded many subtropical and tropical areas of the world including Florida and Hawaii, USA. Laboratory life history and quarantine host range studies of H. memor were conducted with no-choice feeding tests. These tests included eight species of the Anacardiaceae and one species of Sapindaceae. Larvae of H. memor had five to six instars with each head capsule width increasing by 1.68-X. Development time from neonate to adult was 46.7?±?2.2 days. In host range tests, neonates completed development to the adult stage on all non-target species, except Toxicodendron radicans. Moreover, developmental times were delayed and pupal weights were reduced for larvae fed Spondias purpurea leaves. Due to the broad host range exhibited by H. memor larvae, this species will not be considered as a biological control agent of S. terebinthifolia in the continental U.S.A.     

Biology, herbivory, and host specificity of Antiblemma leucocyma (Lepidoptera: Noctuidae) on Miconia calvescens DC. (Melastomataceae) in Brazil

Miconia calvescens DC. (Melastomataceae) is an invasive tree considered one of the greatest threats to natural ecosystems of Hawaii and other Pacific islands. The potential for using the defoliator Antiblemma leucocyma (Lepidoptera: Noctuidae) as a biological control agent of M. calvescens was evaluated in the native habitat of the insect in Brazil. Impact assessment in the field showed that 37.2% of leaves presented damage by A. leucocyma, and among damaged leaves, 1.5-51.9% of the leaf area (16.3% on average) was affected. Damage was found in young to fully expanded leaves (28-915 cm² in area). Individual A. leucocyma developing as third through sixth instars in the laboratory consumed an average of 1348 cm² of M. calvescens leaf material in about 11 days. Populations of A. leucocyma in the field were heavily impacted (up to 83% parasitism) by a braconid parasitoid. Observations in the field and single- and two-host choice tests in the laboratory indicated that A. leucocyma has a narrow host range restricted to M. calvescens. Usefulness of this herbivore for biological control in Hawaii appears low, however, because of its probable susceptibility to generalist parasitoids.     

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.