Seasonal activity,habitat preferences and larval mortality of the leaf-mining fly Calycomyza eupatorivora (Agromyzidae), a biological control agent established on Chromolaena odorata (Asteraceae) in South Africa

The leaf-mining fly Calycomyza eupatorivora Spencer (Diptera: Agromyzidae) has become widely established in the eastern regions of South Africa, following its release for the biological control of the invasive shrub Chromolaena odorata (L.) King and Robinson. This study was conducted to gain some insight into the impact of C. eupatorivora populations, by assessing their seasonal activity, habitat preferences, levels of leaf exploitation and extent of larval mortality in the field over a 1-year period. Leaf mining intensity was poorly synchronised with leaf availability, with leaf exploitation peaking at the end of the growing season of C. odorata. Although significantly more mines were recovered on plants growing in shaded situations, the percentage of available leaves that were exploited for mining was not significantly different between plants growing in shaded versus open situations. Overall, the levels of leaf damage were trivial with mines recovered from <5% of available leaves that were sampled during the study. Besides leaf-quality requirements, these low levels of leaf exploitation may have been influenced by high larval mortality which varied between 60 and 83%, depending on whether lower or higher estimates were used. These evaluations have verified the results of earlier laboratory studies which suggested that the impact of C. eupatorivora on mature populations of C. odorata in South Africa will be negligible.     

Liothrips tractabilis sp.n. (Thysanoptera: Phlaeothripinae) from Argentina, a potential biocontrol agent of Weedy Campuloclinium macrocephalum (Asteraceae) in South Africa

A new species, Liothrips tractabilis, is described from northern Argentina. Feeding by this thrips causes severe damage to the leaves of Campuloclinium macrocephalum, a plant that has been introduced to South Africa where it is a serious weed of grasslands. A key is provided to the four species of the genus Liothrips recorded from Argentina.     

The tortoise beetle Physonota maculiventris (Chrysomelidae: Cassidinae) is suitable for release against the weedy Mexican sunflower Tithonia diversifolia (Asteraceae) in South Africa

The biology and host range of the defoliating beetle Physonota maculiventris Boheman (Chrysomelidae: Cassidinae) were studied in quarantine to determine its suitability for release as a biological control agent against Mexican sunflower, Tithonia diversifolia (Hemsl.) A. Gray (Asteraceae), in South Africa. Females laid 5.25?±?0.25 (mean?±?SE) egg batches during their lifetimes, with each batch consisting of approximately 33 eggs. Larvae were highly gregarious as early instars and both larvae and adults fed voraciously, often defoliating the plants completely. The life cycle of the beetle was completed in 67.5?±?7.5 days under quarantine conditions. Among the 58 test plant species subjected to no-choice tests, P. maculiventris developed successfully on T. diversifolia and some sunflower (Helianthus annuus L.) cultivars. However, only minor damage was recorded on non-target species, notably the exotic weed Xanthium strumarium L. and some H. annuus cultivars. Also, survival to adulthood was considerably lower on sunflower cultivars than on the target weed during these tests. During choice tests, P. maculiventris oviposited and developed successfully on T. diversifolia only, with only minor feeding damage on some H. annuus cultivars, suggesting that the beetle’s field host range will be confined to the target weed. Risk analysis also showed that P. maculiventris presents an extremely low risk to non-target plant species (e.g. within the tribe Heliantheae and other close relatives). The study concluded that P. maculiventris is safe for release in South Africa and an application for its release is being considered by the relevant South African regulatory authorities.     

Molecular relationships between populations of South African citrus thrips (Scirtothrips aurantii Faure) in South Africa and Queensland, Australia

Abstract   In response to the recent establishment of a population of South African citrus thrips ( Scirtothrips aurantii Faure) in Australia, we used DNA sequence data to examine whether this population is distinct from populations in South Africa. Mitochondrial and internal transcribed spacer (ITS2) DNA from populations from different host plants in Australia and South Africa showed no clear separation between populations that was entirely congruent with host plant or country of origin. Analysis of the ITS2 data was confounded by the presence of multiple different copies of the spacer in some populations. Neither the presence or absence, nor the sequence of these copies was clearly diagnostic for any one population. These preliminary data suggest that the Australian population is not a distinct species or subspecies from the populations of S. aurantii on either citrus or Bryophyllum in South Africa.     

Development and characterization of microsatellite markers for the invasive weed Mikania micrantha (Asteraceae)

Mikania micrantha is a successful invasive weed in many parts of the world. Twelve polymorphic microsatellite loci were isolated and characterized from an AC-enriched genomic library of this species. Twenty-eight individuals from one population in Dongguan were tested for polymorphism. The average allele number of these microsatellites was three per locus, ranging from two to five. The expected and observed heterozygosities ranged from 0.275 to 0.742, and from 0.250 to 1.000, respectively. These microsatellite markers can be applied to study the population genetics in the native and invasive ranges of this species, and to trace its invasion history.     

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.     

Inflorescence-inhabiting spiders do not threaten the survival and fecundity of Anthonomus santacruzi (Curculionidae), a florivorous biocontrol agent of Solanum mauritianum (Solanaceae) in South Africa

The florivorous weevil Anthonomus santacruzi is being released in South Africa to reduce the reproductive capacity of the invasive tree, Solanum mauritianum. Initiated to allay concerns about biotic interference, our study suggests that inflorescence-inhabiting spiders pose no risk to the survival and proliferation of A. santacruzi populations in the field.     

Effect of environmental factors on disease development caused by the fungal pathogen Plectosporium alismatis on the floating-leaf stage of starfruit (Damasonium minus), a weed of rice

Starfruit, Damasonium minus (R. Br.) Buch, is a herbicide-resistant, aquatic-weed occurring in Australian rice fields. The fungal pathogen Plectosporium alismatis (Oudem) W.M. Pitt, W. Gams & U. Braun is a potential mycoherbicide for the control of this weed. Previous work showed that the fungus can suppress the growth of starfruit juvenile plants. This paper reports the effect of environmental factors on the disease development on starfruit at the floating-leaf stage. Fungal inoculation of plants in advanced phases of floating leaf stage reduced significantly (P<0.05) the green leaf area, but only when the percentages of diseased areas were deducted. However, inoculation when the first floating leaf was emerging reduced significantly (P<0.001) both, plant dry above-ground biomass and leaf area.     

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.     

The effect of the armored scale,Rhizaspidiotus donacis (Hemiptera: Diaspididae), on shoot growth of the invasive plant Arundo donax (Poaceae: Arundinoideae)

The effect of feeding by the armored scale, Rhizaspidiotus donacis (Leonardi, 1920) (Hemiptera: Diaspididae) on the growth of the plant Arundo donax L. (Poaceae) was evaluated under field conditions in its native range. The study was designed to evaluate the impact of R. donacis, a candidate agent for biological control of A. donax which is invasive in arid riparian ecosystems of the Southwestern USA and Mexico. The study was carried out at five A. donax sites in the Province of Alicante, Spain, differing in altitude and climate. At each site, 30 infested lateral shoots were selected and 15 were randomly treated monthly with imidacloprid insecticide. Shoot lengths were measured monthly over a 1-year period in a comparative growth analysis. Shoots infested with R. donacis had an over 2-fold reduced growth rate as compared to treated shoots. Growth of shoots varied by site, and the effect of R. donacis on growth was most pronounced in the late spring, when mature females produced first instar scale crawlers. The impact of R. donacis on A. donax growth under field conditions in the native range, combined with its narrow host specificity, indicate that R. donacis is a promising candidate for biological control of A. donax in North America and other areas invaded by this weed.     

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.     

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, 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.     

Utilization of an unpredictable food source by Melanterius ventralis, a seed-feeding biological control agent of Acacia longifolia in South Africa

A seed-feeding weevil, Melanterius ventralis (Coleoptera: Curculionidae), has been introduced into South Africa to supplement a gall wasp,Trichilogaster acaciaelongifoliae (Hymenoptera: Pteromalidae), in the biological control programme against an alien invasive tree, Acacia longifolia (Mimosaceae), from Australia. The gall wasp debilitates most of the flower buds on A. longifolia andreduces seed-set by >95%. The intended rolefor M. ventralis is to destroy theresidual seeds. To achieve this, the gravidfemales need to be able to locate a food sourcethat is both heterogeneously dispersed andfrequently scarce due to damage caused by T.acaciaelongifoliae. Observations showedthat M. ventralis females are meticulousin choosing sites to oviposit so that larvae donot become overcrowded and food limited. Cagetests and field observations revealed thatfemales located pods regardless of density andposition, and that the duration of time spenton branches was proportional to the number ofpods on the branches and to the condition ofpods. The females spent little time onbranches of acacia species other than A.longifolia, but some time was spent onbranches with pods of A. melanoxylon thathad been coated with juice extracted from A.longifolia. All indications are that M.ventralis has the attributes needed to bean excellent supplementary biological controlagent to T. acaciaelongifoliae and theprogress of the weevil continues to be monitored.     

Arid-adapted Mexican leafhopper, Barela parvisaccata (Hemiptera: Cicadellidae), unsuitable for biocontrol of Lantana camara (Verbenaceae) in South Africa

A host specificity study conducted in South Africa between 2000 and 2004 showed that the leafhopper, Barela parvisaccata Young, performed as well on some of the indigenous Lippia spp. as on the target weed Lantana camara L. During adult no-choice tests, B. parvisaccata reproduced on eight species of the two closely related genera, Lantana and Lippia. Although B. parvisaccata has a restricted host range, its reproductive performance on the indigenous Lippia scaberrima Sond. during the adult multiple-choice tests was unacceptably high. It was therefore concluded that B. parvisaccata was not sufficiently host-specific to be released against L. camarara in Africa.     

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.     

Effect of temperature on the survival of Aconophora compressa Walker (Hemiptera: Membracidae): implications for weed biocontrol

Abstract  In weed biocontrol, similarity of abiotic factors between the native and introduced range of a biocontrol agent is critical to its establishment and effectiveness. This is particularly the case for weeds that have a wide geographical distribution in the native range. For such weeds, the choice of a specialist insect that has narrow tolerance limits to important abiotic factors can diminish its ability to be an effective biocontrol agent. The membracid Aconophora compressa was introduced in Australia from Mexico for biocontrol of Lantana camara , a plant with a wide climatic tolerance. In this study we investigated the effect of constant and alternating temperatures on A. compressa survival. Longevity of adults and nymphs declined with increasing temperatures, and at 39°C individuals survived for less than a day. At lower temperatures, nymphs survived longer than adults. Survival at alternating temperatures was longer than at constant temperatures, but the general trend of lower survival at higher temperatures remained. Spatially and temporally, the climatic tolerance of A. compressa appears to be a subset of that of lantana, thereby limiting its potential impact.     

Multiple introductions and gene flow in subtropical South American populations of the fireweed,Senecio madagascariensis(Asteraceae)

Non-indigenous plants exhibit different attributes that make them aggressive competitors with indigenous plants and serious threats to biodiversity.Senecio madagascariensis (fireweed, Asteraceae), a native from southern Africa, is a strong competitor in agricultural activities and has toxic alkaloids that may result in high cattle mortality. In Brazil, this weed was collected for the first time in 1995 and has since spread quickly throughout the Pampas region. To better understand the invasion of the fireweed in South America, we used a genetic characterization with internal transcribed spacer (ITS) and microsatellite markers. Based on the ITS data, the southern Brazil populations of S. madagascariensis shared genetic homology with samples taken from the Hawaiian Islands and South Africa. Microsatellite analysis showed the genetic diversity split in two clusters, perhaps intimating the independent introduction of each species into South America. Although fireweed was introduced recently in southern Brazil, the considerable levels of genetic diversity, gene flow, and inbreeding may indicate success in the species establishment in this environment.     

Phylogeography of the invasive weed Hypochaeris radicata (Asteraceae): from Moroccan origin to worldwide introduced populations

In an attempt to delineate the area of origin and migratory expansion of the highly successful invasive weedy species Hypochaeris radicata, we analysed amplified fragment length polymorphisms from samples taken from 44 populations. Population sampling focused on the central and western Mediterranean area, but also included sites from Northern Spain, Western and Central Europe, Southeast Asia and South America. The six primer combinations applied to 213 individuals generated a total of 517 fragments of which 513 (99.2%) were polymorphic. The neighbour-joining tree presented five clusters and these divisions were supported by the results of Bayesian analyses: plants in the Moroccan, Betic Sierras (Southern Spain), and central Mediterranean clusters are all heterocarpic. The north and central Spanish, southwestern Sierra Morena, and Central European, Asian and South American cluster contain both heterocarpic (southwestern Sierra Morena) and homocarpic populations (all other populations). The Doñana cluster includes two homocarpic populations. Analyses of fragment parameters indicate that the oldest populations of H. radicata are located in Morocco and that the species expanded from this area in the Late Quaternary via at least three migratory routes, the earliest of which seems to have been to the southwestern Iberian Peninsula, with subsequent colonizations to the central Mediterranean area and the Betic Sierras. Homocarpic populations originated in the southwestern Iberian Peninsula and subsequently spread across north and central Spain, Central Europe and worldwide, where they became a highly successful weed.