On the origin and genetic variability of the two invasive biotypes of <Emphasis Type="Italic">Chromolaena odorata</Emphasis>

Chromolaena odorata (L.) R. M. King and H. Robinson (Asteraceae), originally from the Neotropics, has become a serious weed in the humid tropics and subtropics of Southeast Asia, Africa and Pacific Islands. In its introduced distributions, C. odorata has been recognised as two biotypes, the Asian/West African (AWA) biotype and South African (SA) biotype, with independent distribution, morphology and ecological characters. To characterise the genetic variability and identify the likely source regions in the native distributions of the two biotypes, we carried out an extensive phylogeographic study using chloroplast and nuclear DNA sequences and microsatellite DNA markers. The analysis of both DNA sequences and nuclear markers showed that native populations possessed high genetic diversity, while both the AWA and SA biotypes in invaded regions appeared to have low genetic diversity. The AWA and SA biotypes were genetically distinct. Strong competitive ability and environmental adaptability may have facilitated the invasion AWA and SA biotypes in its respective invasive regions. We conclude that the source of AWA biotype may be Trinidad and Tobago, while the SA biotype was from Cuba and Jamaica. For a better outcome of biocontrol, the potential biological control agents for the two biotypes should be collected from these native regions, respectively.     

First record of a specialist folivore of Chromolaena odorata (Asteraceae) in Togo,and indices of its range expansion in Nigeria: implications for biological control

Pareuchaetes pseudoinsulata is reported for the first time in Togo and widespread in south-western Nigeria, albeit in low densities and difficult to detect. Country-wide surveys are warranted in both countries, with the intention of renewing efforts in the biological control of Chromolaena odorata in western Africa, where the socio-ecological impact of the weed is significant.     

Host specificity ofPareuchaetes insulata (Lep.: Arctiidae), a biological control agent forChromolaena odorata (Compositae)

Larvae of the arctiid moth,Pareuchaetes insulata, from Florida fed on the leaves ofChromolaena odorata, a serious composite, alien weed in Natal, South Africa. In starvation test trials using 48 plant species,P. insulata completed its development onC. odorata andAgeratum houstonianum. Subsequent attempts to cultureP. insulata on these two plants was only successful onC. odorata. The biology ofP. insulata is similar to that of two other well studiedPareuchaetes species namelyP. pseudoinsulata andP. aurata aurata. Repeated defoliation ofC. odorata byP. insulata could contribute to its control as has been found withP. pseudoinsulata in Guam.Pareuchaetes insulata has been approved for release as a biological control agent ofC. odorata in South Africa.     

The host specificity and climatic suitability of the gall fly Cecidochares connexa (Diptera: Tephritidae), a potential biological control agent for Chromolaena odorata (Asteraceae) in Australia

The gall fly Cecidochares connexa (Diptera: Tephritidae) is a potential biological control agent for Chromolaena odorata in Australia. Its host specificity was determined against 18 species in the tribe Eupatorieae (Family Asteraceae) in which C. odorata belongs, in quarantine in Brisbane, Australia. Oviposition occurred and flies developed on only C. odorata and Praxelis clematidea, both of which are in the subtribe Praxelinae. P. clematidea is considered a weed outside tropical America. In both multiple-species-minus-C. odorata choice tests and single-species no-choice tests, the mean number of galls/plant was significantly greater on C. odorata (48 and 41, respectively) than on P. clematidea (2 and 9, respectively). There were also significantly more adults emerging from C. odorata (mean 129 and 169, respectively) in the two types of tests than from P. clematidea (1 and 8, respectively). Paired choice, multiple generation (continuation) and time dependent tests further clarified the extent that C. connexa could develop on P. clematidea. In these tests, the mean number of galls formed and the mean number of emerging adults were consistently less for P. clematidea than C. odorata and populations of C. connexa could not be maintained on P. clematidea. Galls were not seen on any other plant species tested. This study supports the results of host specificity testing conducted in seven other countries and confirms that C. connexa poses little risk to other plant species in Australia. C. connexa has been released in 10 countries and an application seeking approval to release in Australia has been submitted to the Australian Government.     

Impact of the shoot-boring moth Dichrorampha odorata (Lepidoptera: Tortricidae) on growth and reproductive potential of Chromolaena odorata (Asteraceae) in the laboratory

A 9-month laboratory study was carried out to determine the impact of herbivory by a moth with shoot-boring larvae, Dichrorampha odorata Brown and Zachariades (Lepidoptera: Tortricidae) on growth and reproductive ability of its host plant, Chromolaena odorata (L.) King and Robinson (Asteraceae), a major invasive alien plant species in southern Africa. Newly hatched D. odorata larvae were inoculated onto 0 (control), 50 and 100% of the shoot tips of C. odorata in the laboratory. At all treatment levels, the basal stem diameter of C. odorata was not affected by D. odorata larval feeding. Larval feeding by D. odorata significantly reduced the height of the main shoot and flower production in C. odorata relative to the control treatment but promoted branching by increasing the number of shoot tips. However, the differences in plant height and number of flowers between the 50 and 100% inoculation levels were not statistically significant. Dichrorampha odorata is the first shoot-tip attacking agent that is being released as a biological control agent against C. odorata in South Africa. In general, the impacts of D. odorata on the weed were relatively small even though statistically significant. The findings of this study suggest that high levels of damage by the moth will modestly reduce the height, flower production, and the competitiveness of C. odorata, thereby contributing to the biological control of the weed in South Africa.     

Prospects for the classical biological control of Calotropis procera (Apocynaceae) using coevolved insects

Calotropis procera (Apocynaceae), a native of tropical Africa, the Middle East and the Indian subcontinent, is a serious environmental and rangeland weed of Australia and Brazil. It is also a weed in Hawaii in the USA, the Caribbean Islands, the Seychelles, Mexico, Thailand, Vietnam and many Pacific Islands. In the native range C. procera has many natural enemies, thus classical biological control could be the most cost-effective option for its long-term management. Based on field surveys in India and a literature search, some 65 species of insects and 5 species of mites have been documented on C. procera and another congeneric-invador C. gigantea in the native range. All the leaf-feeding and stem-boring agents recorded on Calotropis spp. have wide host range. Three pre-dispersal seed predators, the Aak weevil Paramecops farinosus, the Aak fruit fly Dacus persicus in the Indian subcontinent and the Sodom apple fruit fly Dacus longistylus in the Middle East, have been identified as prospective biological control agents based on their field host range. In Australia and Brazil, where C. procera has the potential to spread across vast areas, pre-dispersal seed predators would help to limit the spread of the weed. While the fruits of C. procera vary in size and shape across its range, those from India are similar to the ones in Australia and Brazil. Hence, seed-feeding insects from India are more likely to be suitable due to adaptation to fruit size and morphology. Future survey efforts for potential biological control agents should focus on North Africa.     

Biological control of the weed,Chromolaena odorata [Asteraceae], byPareuchaetes pseudoinsulata [Lep.: Arctiidae] on Guam and the northern Mariana Islands

Chromolaena odorata L. is a major weed of tropical plantations and pasture lands.Pareuchaetes pseudoinsulata Rego Barros [Lep.: Arctiidae] was mass-reared and released to control the weed on the Mariana Islands of Guam, Rota, Tinian and Saipan in the western Pacific. One year following establishment ofP. pseudoinsulata on Guam, the weed was reduced 100% at 3 of 4 sites monitored with fixed quadrats. Seed production was prevented if defoliation occurred prior to the beginning of the flowering season. If defoliation occurred shortly after flowering began, the developing seeds were eaten byP. pseudoinsulata or there was a significant reduction in weight and germination of seeds that matured. Eighteen months after establishing on Guam,P. pseudoinsulata had defoliated approximately 25,000 hectares of the weed. This is the 1^st report on the quantitative impact of any control agent forC. odorata, and though the long term effect of the insect is still unknown, these results are encouraging.     

Parasitoids of the arctiid mothPareuchaetes pseudoinsulata (Lep.: Arctiidae), an introduced biocontrol agent against the weedChromolaena odorata (asteraceae), in asia and Africa

The leaf-feeding arctiid mothPareuchaetes pseudoinsulata has been released widely in tropical countries for the control of the weedChromolaena odorata. P. pseudoinsulata is now widespread and locally abundant in Ghana, Indonesia, India, Sri Lanka, Malaysia, Micronesia and the Philippines. In the West Indies where it is native,P. pseudoinsulata is host to 2 species of egg parasitoids, 4 species of tachinid larval parasitoids, 1 larval-pupal parasitoid, and 1 pupal parasitoid, as well as a virulent nuclear-polyhedrosis virus.P. pseudoinsulata is not significantly parasitised where it is established in south-east Asia, Oceania or Africa. No egg parasitoids have been reported, and only 3 larval or larval-pupal parasitoids. Only one of these, the larval-pupal tachinidExorista xanthaspis, is regularly encountered, and rarely exceeds 10% parasitism in the field. These results confirm the pattern that herbivorous insects from families with exposed larvae are unlikely to be heavily parasitised in a new geographic range.     

The nocturnal larvae of a specialist folivore prefer <Emphasis Type="Italic">Chromolaena odorata</Emphasis> (L.) foliage from a sunny environment,but does it matter?

Increasing evidence suggests that the responses of insect herbivores to environment-mediated changes in the phenotypic and phytochemical traits of their host plants are more complex than previously thought. Here, we examined the effects of habitat conditions (shaded versus full-sun habitats) on plant traits and leaf characteristics of the invasive alien plant, Chromolaena odorata (L.) (Asteraceae). We also determined neonate larval preference of the specialist herbivore, Pareuchaetes pseudoinsulata Rego Barros (Lepidoptera: Erebidae) (a biological control agent) for shaded versus full-sun leaves. The study further evaluated the performance of the moth on C. odorata leaves obtained from both shaded and full-sun habitats. Leaves of C. odorata plants growing in the shaded habitat had higher water and nitrogen contents compared with full-sun leaves. Plants growing in shade had longer leaves but full-sun plants were taller and had greater aboveground biomass compared with shaded plants. Although neonate larvae of P. pseudoinsulata preferred to feed on full-sun foliage, development was faster when reared on shaded foliage. However, survival, pupal mass, growth rate, and Maw’s host suitability index of the moth did not significantly differ between full-sun and shaded foliage. Our inability to demonstrate significant differences in key insect performance metrics in P. pseudoinsulata between shaded and full-sun foliage, despite neonate larval preference for one of the foliage types, suggests that neither of the foliage types can be considered a superior host, and reiterate the fact that relationships between host plant quality (modulated by light intensity) and phytophagous insect performance are not simple.     

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.     

Interactions between Chromolaena odorata (Asteraceae) and Pareuchaetes pseudoinsulata (Lepidoptera: Arctiidae)

Feeding of Pareuchaetes pseudoinsulata caterpillars caused the leaves of Chromolaena odorata to turn yellow. Leaf yellowing could not be induced either by artificial removal of leaves or by drenching the plant with a solution of excreta from P. pseudoinsulata caterpillars. Yellow leaves appeared tougher but had the same energy level as that of green leaves. The amount of nitrate-nitrogen was significantly higher in yellow leaves than green leaves. P. pseudoinsulata caterpillars prefer to feed on green leaves. When forced to feed on yellow leaves, they exhibit slow growth and high mortality. Defensive factors in plants attacked by insects seemed to prevent further infestation of plants. In the field, caterpillars on the yellow plants were found during both day and night whereas on green plants they appeared to feed at night and hide in the ground at daytime.     

Variation in host plant has no effect on the performance and fitness‐related traits of the specialist herbivore Pareuchaetes insulata

The effect of host plant dissimilarity on insect preference and performance was tested using two morphological forms of Chromolaena odorata (L.) King & Robinson (Asteraceae) (one from Florida, USA, another from South Africa), and a specialist herbivore, Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae: Arctiinae) from Florida, USA, that was introduced as a biological control agent in South Africa. Although this insect did establish at one site, of some 21 sites at which over 800 000 individuals were released, its population level in the field has remained low after an initial outbreak in 2006. To explain the poor performance of P. insulata, we hypothesised that P. insulata larvae prefer Floridian C. odorata to the southern African C. odorata, which is morphologically and probably genetically distinct, and that larvae reared on Floridian C. odorata should have higher fitness and performance. We tested this by comparing insect performance metrics on each of the two plant forms in laboratory experiments. Apart from pupal mass, which was significantly greater on southern African C. odorata, P. insulata performance metrics were similar on both plant forms; there were no significant differences in total leaf area consumed, egg and larval development, immature survival rates, feeding index, host suitability index, growth index, and fecundity between the Floridian and southern African C. odorata plants. In sum, we could not demonstrate that differences in plant forms in C. odorata are responsible for the poor performance of P. insulata in South Africa.     

Phytoremediation of fuel oil and lead co-contaminated soil by Chromolaena odorata in association with Micrococcus luteus

Phytoremediation is widely promoted as a cost-effective technology for treating heavy metal and total petroleum hydrocarbon (TPH) co-contaminated soil. This study investigated the concurrent removal of TPHs and Pb in co-contaminated soil (27,000 mg kg^?1 TPHs, 780 mg kg^?1 Pb) by growing Siam weed (Chromolaena odorata) in a pot experiment for 90 days. There were four treatments: co-contaminated soil; co-contaminated soil with C. odorata only; co-contaminated soil with C. odorata and Micrococcus luteus inoculum; and co-contaminated soil with M. luteus only. C. odorata survived and grew well in the co-contaminated soil. C. odorata with M. luteus showed the highest Pb accumulation (513.7 mg kg^?1) and uptake (7.7 mg plant^?1), and the highest reduction percentage of TPHs (52.2%). The higher TPH degradation in vegetated soils indicated the interaction between the rhizosphere microorganisms and plants. The results suggested that C. odorata together with M. luteus and other rhizosphere microorganisms is a promising candidate for the removal of Pb and TPHs in co-contaminated soils.     

Invasion genetics of Chromolaena odorata (Asteraceae): extremely low diversity across Asia

Chromolaena odorata is a native of America while a weed in many parts of tropical and subtropical regions in the world. Research into the invasion mechanisms of C. odorata contributes to a broader understanding of factors that facilitate plant adaptation, and also helps developing effective management strategies. In this study, we used three DNA fragments and six microsatellite loci: (1) to compare genetic diversity of C. odorata in its native and invaded regions; (2) to elucidate the invasive routes and identify possible source locations of C. odorata from America to Asia, with attempt to evaluate the possible mechanisms facilitating the successful invasion of this species. Despite two recorded independent introductions, DNA sequence data revealed only one single haplotype of C. odorata present throughout tropical Asia. All six microsatellite loci consistently exhibited extremely low genetic diversity in Asian populations compared to those from native ranges. Our results implied that there was likely only a single introduction to Asia, and Trinidad, Tobago and adjacent areas in the West Indies were the most likely source location of that introduction. The successful invasion of C. odorata in Asia may have been facilitated by the genotype with strong competitive ability.     

<Emphasis Type="Italic">Chromolaena odorata</Emphasis> (Siam weed) in eastern Africa: distribution and socio-ecological impacts

Invasive alien plant species such as Chromolaena odorata have negative impacts on biodiversity, ecosystem services and human well-being. Ecological impacts of this shrub are relatively well understood, but its impacts on local livelihoods and perceptions are poorly documented. We mapped C. odorata distribution in eastern Africa (Ethiopia, Kenya, Rwanda, Tanzania and Uganda) and compared perceptions and quantified the impacts of this species across Tanzanian villages with varying degrees of invasion density. Data were collected through 240 household questionnaires. Results indicate that C. odorata is a relatively new invader that already has severe negative impacts and is threatening livelihoods and the environment. Impacts include reductions in native biodiversity and the amount of available forage for livestock, reduced crop and water yields, and impaired mobility. Continued spread will cause additional negative impacts on poor rural communities. Implementation of a biological control programme targeting C. odorata is needed as a cost effective management approach along with other control and restoration measures.     

Zooplankton characteristics in monitoring of Lake Ladoga

Water lettuce (Pistia stratiotes L. (Araceae)) is alien to Africa and a declared weed in South Africa. In large perennial rivers it is effectively controlled by its biological control agent, Neohydronomus affinis (Coleoptera: Curculionidae). On those shallow isolated water bodies which are regularly subjected to alternate wet and dry regimes that become infested with water lettuce, chemical control is necessary to prevent further spread of the weed and to facilitate access to water. This paper discusses the short-term chemical control and the long-term biological control of water lettuce. The need for further research is outlined.     

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.     

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.     

Field assessment of a frond-feeding weevil, a successful biological control agent of red waterfern, Azolla filiculoides, in southern Africa

Azolla filiculoides (red waterfern) is a small, floating fern native to South America, that has invaded aquatic habitats, predominantly water resevoirs in southern Africa. A frond-feeding weevil, Stenopelmus rufinasus Gyllenhal (Coleoptera: Curculionidae), was imported from Florida, USA, and released as a biological control agent against this weed in South Africa at the end of 1997. To date, 24,700 weevils have been released, which has resulted in local extinction of red waterfern at 81% of the 112 release sites. The weevil has not failed to control a single site. Several sites were, however, lost due to flooding or drainage of dams. The surface area of weed controlled totalled 203.5 ha. On average, A. filiculoides was controlled in infested sites in 6.9 (±4.3) months. The weed recolonized at 22 of the sites (through either spore germination or dispersal by waterfowl), but the weevils subsequently spread to all of these sites and successfully caused local extinction of the weed at 18 of the sites. Five years after the release of the weevil, the weed no longer poses a threat to aquatic systems in southern Africa. In comparison to other biological control programs of aquatic weeds, the program against A. filiculoides in southern Africa ranks among the most successful cases anywhere in the world.     

Tri-Trophic Insecticidal Effects of African Plants against Cabbage Pests

Botanical insecticides are increasingly attracting research attention as they offer novel modes of action that may provide effective control of pests that have already developed resistance to conventional insecticides. They potentially offer cost-effective pest control to smallholder farmers in developing countries if highly active extracts can be prepared simply from readily available plants. Field cage and open field experiments were conducted to evaluate the insecticidal potential of nine common Ghanaian plants: goat weed, Ageratum conyzoides (Asteraceae), Siam weed, Chromolaena odorata (Asteraceae), Cinderella weed, Synedrella nodiflora (Asteraceae), chili pepper, Capsicum frutescens (Solanaceae), tobacco, Nicotiana tabacum (Solanaceae) cassia, Cassia sophera (Leguminosae), physic nut, Jatropha curcas (Euphorbiaceae), castor oil plant, Ricinus communis (Euphorbiaceae) and basil, Ocimum gratissimum (Lamiaceae). In field cage experiments, simple detergent and water extracts of all botanical treatments gave control of cabbage aphid, Brevicoryne brassicae and diamondback moth, Plutella xylostella, equivalent to the synthetic insecticide Attack^® (emamectin benzoate) and superior to water or detergent solution. In open field experiments in the major and minor rainy seasons using a sub-set of plant extracts (A. conyzoides, C. odorata, S. nodiflora, N. tabacum and R. communis), all controlled B. brassicae and P. xylostella more effectively than water control and comparably with or better than Attack^®. Botanical and water control treatments were more benign to third trophic level predators than Attack^®. Effects cascaded to the first trophic level with all botanical treatments giving cabbage head weights, comparable to Attack^® in the minor season. In the major season, R. communis and A conyzoides treatment gave lower head yields than Attack^® but the remaining botanicals were equivalent or superior to this synthetic insecticide. Simply-prepared extracts from readily-available Ghanaian plants give beneficial, tri-trophic benefits and merit further research as an inexpensive plant protection strategy for smallholder farmers in West Africa.