Prioritisation of potential agents for the biological control of the invasive alien weed,Pereskia aculeata (Cactaceae), in South Africa

Pereskia aculeata Miller (Cactaceae) is an invasive alien species in South Africa that is native in Central and South America. In South Africa, P. aculeata outcompetes native plant species leading to a reduction in biodiversity at infested sites. Herbicidal and mechanical control of the plant is ineffective and unsustainable, so biological control is considered the only potential solution. Climatic matching and genotype matching indicated that the most appropriate regions in which to collect biological control agents were Santa Catarina and Rio de Janeiro provinces in Southern Brazil. Surveys throughout the native distribution resulted in 15 natural enemy species that were associated with the plant. Field host range data, as well as previous host plant records, were used to prioritise which of the species were most likely to be suitably host specific for release in South Africa. The mode of damage was used to determine which species were most likely to be damaging and effective if released. The most promising species prioritised for further study, including host specificity and impact studies, were the stem-wilter Catorhintha schaffneri Brailovsky & Garcia (Coreidae); the stem boring species Acanthodoxus machacalis Martins & Monné (Cerambycidae), Cryptorhynchus sp. (Curculionidae) and Maracayia chlorisalis (Walker) (Crambidae) and the fruit galler Asphondylia sp. (Cecidomyiidae). By prioritising the potential biological control agents that are most likely to be host-specific and damaging, the risk of conducting host specificity testing on unsuitable or ineffective biological control agents is reduced.     

Pereskiophaga brasiliensis,a natural enemy of the invasive alien cactus Pereskia aculeata,is not suitably host specific for biological control in South Africa

The stem-mining weevil, Pereskiophaga brasiliensis, was a candidate biological control agent for the invasive cactus Pereskia aculeata in South Africa. In host specificity trials, it developed on two indigenous test plant species under choice and no-choice conditions. Pereskiophaga brasiliensis is therefore not suitably host specific for release 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.     

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.     

Biology of Polypedilum n. sp. (Diptera: Chironomidae), a promising candidate agent for the biological control of the aquatic weed Lagarosiphon major (Hydrocharitaceae) in Ireland

Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae), an aquatic macrophyte native to Southern Africa that has become invasive in several countries worldwide, is a potential target for biological control. Biology studies were conducted on Polypedilum (Pentapedilum) n. sp. near reei Oyewo & Sæther (Diptera: Chironomidae), a midge whose larvae were discovered mining the plant's shoot tips in its native range. Field surveys indicated that the midge occurred only at a small number of sites but attained high densities (up to 370 shoots damaged/m²) that prevented further growth from the shoot tips. A population of the midge was imported into quarantine in Ireland to evaluate its potential as a candidate biological control agent. The adult stage is terrestrial and short-lived (4–5 d), with females depositing one–two egg packets into the water bodies. First-instar larvae fed externally on the stems and leaflets. Later instars fed on the apical meristems and burrowed into the shoot tips, with resultant damage stunting the apical growth. Larvae moved readily between shoots to complete their development and pupated within the tunnels excavated by the late instar larvae. Developmental time to adulthood ranged from 31 to 49 days at 20.7°C and a 15 L:9-D cycle. This is the first time that a chironomid has been imported and successfully cultured for consideration as a classical biological control agent. Several aspects of the midge's biology suggest that host specificity testing is warranted to determine its potential as a biocontrol agent of L. major.     

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, host specificity and risk assessment ofGargaphia decoris, the first agent to bereleased in South Africa for the biological control ofthe invasive tree Solanum mauritianum

The South American tree Solanummauritianum is a major environmental weed in thehigh-rainfall regions of South Africa and has beentargeted for biological control since 1984. Althoughhost ranges of imported agents determined duringquarantine tests have resulted in the rejection ofeight of the 11 candidate agents tested so far, theleaf-sucking lace bug Gargaphia decoris,imported from Argentina in 1995, displayed anacceptably narrow host range in captivity. No-choicetests showed that G. decoris is confined to Solanum species and cannot survive on solanaceouscrops outside that genus. Although these tests alsoindicated that G. decoris colonies could surviveand reproduce on cultivated eggplant (aubergine) andat least five native South African Solanumspecies, all but one native species proved to beinferior hosts in terms of adult survival andoviposition potential. During paired choice andmultichoice tests in small and larger cagesrespectively, G. decoris displayed very strongfeeding and oviposition preferences for S. mauritianum. Analyses of the risk of attack onnon-target Solanum plants revealed that, withone possible exception, none were likely to suffermore than incidental damage in the field. Host recordsfrom South America have also indicated that G. decoris has not been recorded on any Solanumspecies other than S. mauritianum, providingfurther evidence of its host specificity. The resultsof this study were accepted by the regulatoryauthorities and in February 1999, G. decorisbecame the first agent to be released in South Africafor the biological control of S. mauritianum.     

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.     

Biology and host range of Mada polluta,a potential biological control agent of Tecoma stans in South Africa

The host range of Mada polluta Mulsant (Coleoptera:Coccinnellidae) was studied to assess its suitability as a biological control agent of Tecoma stans (L.) Juss ex Kunth var stans (Bignoniaceae), an invasive weed in South Africa. Biology of M. polluta and its host range were determined in the laboratory using no-choice and multi-choice feeding, oviposition and larval survival tests. Out of 36 plant species from 12 plant families (Bignoniaceae, Acanthaceae, Asteraceae, Verbenaceae, Lamiaceae, Oleaceae, Cucurbitaceae, Fabaceae, Scrophulariaceae, Solanaceae, Apiaceae, Chenopodiaceae and Poaceae) within the order Lamiales that were tested during the host specificity testing, M. polluta showed a very strong preference for T. stans, depositing its eggs on T. stans and none on non-target plant species. In no-choice tests, an average of 246 eggs was laid on T. stans, and from these, 133 larvae developed to adulthood. The beetle also showed very promising biological attributes that will contribute to its success as a biological control agent of T. stans. These attributes include highly damaging larvae and adults, high fecundity (532 eggs/female) and a short life cycle (four weeks). The short life cycle will enable multiple generations per year and rapid population increase in the field. It is concluded that M. polluta is sufficiently host-specific to be released against T. stans in South Africa. It is, therefore, strongly recommended that permission be granted to release this beetle from quarantine for biological control of T. stans in South Africa.     

Should the mirid,Eccritotarsus catarinensis (Heteroptera: Miridae), be considered for release against water hyacinth in the United States of America?

Between one and seven biological control agents have been released against water hyacinth (Eichhornia crassipes (Mart.) Solms) in at least 30 countries, with varied success. A mirid, Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae), the most recent agent released, is damaging to the plant on the African continent. It could be useful in the USA where water hyacinth remains a problem, but its introduction remains in doubt because during host specificity trials, it developed on Pontederia cordata L. (pickerelweed), indigenous to the USA. However, it did not establish on pickerelweed monocultures during South African field trials, and only light spillover feeding occurred where the two plants coexisted suggesting that the use of P. cordata as a host is a laboratory artefact and it may be suitable for use in the USA, if its thermal physiology allows establishment. We reran models developed for South Africa using CLIMEX to predict whether the mirid will establish where water hyacinth and pickerelweed co-occur, but not where pickerelweed occurs in the absence of water hyacinth. The models suggest that the mirid's distribution will be limited by cold winter temperatures and insufficient thermal accumulation to the southern states of the USA, within the main distribution of water hyacinth. Even though some spillover feeding on pickerelweed might result where the two plants co-occur, the risk of population level effects seems minimal and the risk to more northern pickerelweed negligible. The benefits, including improved habitat for pickerelweed, associated with further suppression of water hyacinth, outweigh the minimal risk of collateral damage to pickerelweed.     

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.     

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.     

The potential for birds to disperse the seeds of Acacia cyclops,an invasive alien plant in South Africa

Rooikrans Acacia cyclops is an aggressive invasive tree that threatens natural resources in South Africa. The seeds of A. cyclops have a prominent aril which attracts birds that ingest the seeds and disperse them endozoochorously. Two biological control agents, a Seed Weevil Melanterius servulus and a Flower‐galling Midge Dasineura dielsi, were released on A. cyclops in 1991 and 2002, respectively. Together these agents have substantially reduced seed production and generally far lower numbers of seeds are now available to birds. A consequence of this transition from historically bounteous quantities of seeds to scanty seed availability is that birds may no longer associate with the trees and seed dispersal may be disproportionately reduced. To assess whether this has happened, seed attrition was measured by comparing the amount of seeds that disappeared from two groups of branches, one available to birds and the other enclosed in bird netting. Other types of granivores (mainly field mice) were excluded from both groups of branches with a plastic funnel placed around the stems. Mature seeds were also harvested and fed to caged bird species to determine gut retention times and germination rates of ingested seeds. Attrition rates of seeds showed that birds continue to remove seeds but that only a proportion of the crop is taken. Only two frugivorous species (Knysna Turaco Tauraco corythaix and Red‐winged Starling Onychognathus morio) and two granivorous species (Red‐eyed Dove Streptopelia semitorquata and Laughing Dove Streptopelia sengalensis) ingested A. cyclops seeds during feeding trials. Ingestion by birds enhanced seed germination except for those ingested by Laughing Doves. There were no apparent effects of length of gut passage time and avian body size on seed germination rates. Despite the diminished seed resource due to biological control agents, birds continue to disperse A. cyclops seeds.     

Biology, host specificity and risk assessment of the leaf-mining flea beetle, Acallepitrix sp. nov., a candidate agent for the biological control of the invasive tree Solanum mauritianum in South Africa

The South American tree Solanum mauritianum, a major environmentalweed in the high-rainfall regions of SouthAfrica, has proved to be a difficult target forbiological control. Artificially expanded hostranges of candidate agents during quarantinehost-specificity tests, have resulted in therelease of only one agent species which has sofar had a negligible impact on the weed. Thenecessity for additional agents resulted in theimportation of a leaf-mining flea beetle, anunidentified species of Acallepitrix,from Brazil in 1997. No-choice tests inquarantine showed that potential host plantsare confined to the genus Solanum.Although several non-target plants, includingcultivated eggplant (aubergine) and some nativeSouth African Solanum species, sustainedfeeding, oviposition and the development oflarval leaf mines, most of these were inferiorhosts. These results were confirmed by pairedchoice tests, where Acallepitrix sp. nov.displayed significant feeding and ovipositionpreferences for S. mauritianum and where,with few exceptions, more larval leaf mineswere initiated on S. mauritianum.Interpretation of the host-specificity testswas facilitated by a risk assessment matrixwhich suggested that the risk of feeding andoviposition on non-target Solanum plants,including eggplant cultivations, was relativelylow. These and other considerations, such asthe lack of evidence of damage inflicted oncultivated Solanaceae in South America, suggestthat Acallepitrix sp. nov. could beconsidered for release against S. mauritianum in South Africa. However, the results of the host-specificity tests remain ambiguous and until more compelling evidence is obtained from field surveys and open-field trials in Brazil, an application for permission to release the flea beetle in South Africa will not be considered.     

The thermal physiology of Stenopelmus rufinasus and Neohydronomus affinis (Coleoptera: Curculionidae), two biological control agents for the invasive alien aquatic weeds,Azolla filiculoides and Pistia stratiotes in South Africa

Water lettuce, Pistia stratiotes, and red water fern, Azolla filiculoides, are floating aquatic macrophytes that have become problematic in South Africa. Two weevils, Neohydronomus affinis and Stenopelmus rufinasus, are successful biological control agents of these two species in South Africa. The aim of this study was to investigate the thermal requirements of these two species to explain their establishment patterns in the field. Laboratory results showed that both weevils are widely tolerant to cold and warm temperatures. The critical thermal minima (CT_min) of N. affinis was determined to be 5.58?±?0.31°C and the critical thermal maxima (CT_max) was 44.52?±?0.27°C, while the CT_min of S. rufinasus was 5.38?±?0.33°C and the CT_max?44.0?±?0.17°C. In addition, the lower lethal temperatures were ?9.85?±?0.06°C for N. affinis and ?6.85?±?0.13°C for S. rufinasus, and the upper lethal temperatures were 42.7?±?0.85°C for N. affinis and 41.9?±?2.52°C S. rufinasus. Using the reduced major axis regression method, the development for N. affinis was described using the formula y?=?12.976x?+?435.24, while the development of S. rufinasus was described by y?=?13.6x?+?222.45. These results showed that S. rufinasus develops twice as fast as N. affinis. Using these formulae and temperature data obtained from the South African Weather Service, N. affinis was predicted to complete between 4 and 9 generations per year in South Africa, while S. rufinasus was predicted to complete between 5 and 14 generations per year around the country. These results suggest that both species should not be limited by cold winter, nor warm summer temperatures, and should establish throughout the ranges of the weeds in South Africa.     

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

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

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.     

Using molecular methods to determine the origin of weed populations of Pereskia aculeata in South Africa and its relevance to biological control

New biological control agents are required in order to reach and sustain an adequate level of control of the declared environmental weed Pereskia aculeata Miller (Cactaceae) in South Africa. Identifying the origin of weed genotypes has been important in a number of biological control programmes and is likely to be of importance for the control of P. aculeata due to its disjunct native distribution and morphological polymorphisms between plants from different regions of the native and introduced distribution. DNA sequencing of the trnL chloroplastic intron and the phyC nuclear gene indicate that the South African weed population’s origin was in the southern region of native distribution. Inter-Simple Sequence Repeats (ISSRs) confirmed this result and added resolution to the analysis indicating that the native plants with the closest genetic distance to the South African weed population were found in Rio de Janeiro Province, Brazil. The relationship between the South African weed population and garden variety plants as well as the large genetic distance between the South African plants and the native plants suggests that the South African population may be the progeny of escaped garden variety plants that have been cultivated and possibly hybridized. The low levels of genetic variation within the South African population and the monophyly of the South African plants indicates that these plants are the progeny of a single introduction or multiple introductions from the same source. Rio de Janeiro Province in Brazil is the most appropriate region in which to survey for new biological control agents.     

Megamelus scutellaris (Hemiptera: Delphacidae), a biocontrol agent of water hyacinth,is not sufficiently specific for release in Australia

Water hyacinth Eichhornia crassipes (Pontederiaceae) is one of the world's worst invasive species, responsible for damaging aquatic systems in many warmer parts of the globe including north America, Africa, Asia and Australia. The planthopper Megamelus scutellaris Berg (Delphacidae) has been released in USA and approved for release in South Africa for biocontrol of water hyacinth. We assessed this agent for suitability for release in Australia and found that a related native aquatic plant, Monochoria cyanea (Pontederiaceae) is within the fundamental host range of this insect. Adult survival, oviposition and development of nymphs to adult was equally high on M. cyanea as on the target species, although the quality of these next generation adults was lower than those reared on the target species. This demonstrates that M. scutellaris is not sufficiently specific for release in Australia. Nymphal development to adults occurred only in very low numbers on the three other Australian species of Monochoria. M. cyanea only occurs in Australia so M. scutellaris is still a possible water hyacinth biocontrol candidate for other regions depending on the results of assessment of the risk to local species of Monochoria. This study demonstrates the effectiveness of modern biocontrol agent assessment and reinforces the importance of testing of local non-target species.