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.     

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.     

Eight decades of invasion by Chromolaena odorata (Asteraceae) and its biological control in West Africa: the story so far

ABSTRACT

Chromolaena odorata (L.) R.M. King and H. Robinson (Asteraceae) is a perennial weedy shrub of neotropical origin and a serious biotic threat in its invasive range. The Asian-West Africa (AWA) biotype of C. odorata present in West Africa is both morphologically and genetically different from the southern African (SA) biotype. The AWA biotype was first introduced into Nigeria in the late 1930s and rapidly spread across West Africa. Currently, 12 of the 16 countries in West Africa have been invaded, with significant negative effects on indigenous flora and fauna. However, locals in West Africa have found several uses for the weed. As chemical, physical and other conventional methods were unsustainable, costly and largely ineffective, three biological control agents, Apion brunneonigrum (Coleoptera: Brentidae), Pareuchaetes pseudoinsulata (Lepidoptera: Erebidae) and Cecidochares connexa (Diptera: Tephritidae), have been released in West Africa between the 1970s and the early 2000s. However, only C. connexa and P. pseudoinsulata established, contributing to the control of the weed, in six and four countries in West Africa respectively. Limited research funding, the absence of post-release evaluations of the established agents, and the ‘conflict of interest’ status of C. odorata (i.e. being beneficial for local use but damaging to ecosystem services and agriculture), are serious factors deterring the overall biological control effort. Here, using historical records and field surveys, we examine the invasion history, spread, impacts, and management of C. odorata in West Africa and make recommendations for the sustainable management of C. odorata in the region.     

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.     

Nitrogen fertilisation improves growth of Chromolaena odorata (Asteraceae) and the performance of the biological control agent,Pareuchaetes insulata (Erebidae)

Recent studies have demonstrated, through their contrasting results, that relationships between nitrogen levels in host plants and phytophagous insect performance are not simple. This study examined the effect of varying fertilisation regimes on the invasive alien plant, Chromolaena odorata (L.) (Asteraceae) and the response of a specialist folivore (a biological control agent), Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae). C. odorata plants were treated with 3 different levels of fertilisation and plant characteristics were measured within 2–3 months of fertiliser application. Leaves from each of the three treatments were fed to newly hatched larvae until pupation in order to determine the effect of nitrogen fertilisation on herbivore performance metrics such as survival, development time, fecundity and longevity. High and medium fertilisation significantly increased foliar nitrogen concentrations, basal stem diameter, leaf length, shoot height and above-ground biomass of C. odorata plants relative to low fertilisation. When individuals of P. insulata were fed on leaves from medium- or high-fertilisation treatments, they developed faster, grew to a larger size (by 8%) and achieved higher fecundity (19–22%) than leaves from the low-fertilisation treatment. The results suggest that in mass-rearing, increased production of this biological control agent will occur in high- or medium-fertilised plants.     

两种本地植物种子萌发对飞机草的化感耐受性及其相互竞争

为探讨本地物种假地豆和白饭树对入侵植物飞机草的替代控制潜力,利用培养皿法和同质园种植实验分别研究了两个本地物种种子萌发对飞机草的化感耐受性及其与飞机草的竞争关系。结果显示:除了假地豆的萌发率在高浓度(2.5%)的飞机草叶提取液下受到显著抑制外,两个本地种的萌发在不同浓度飞机草根、茎、叶提取液下均不受抑制。飞机草与假地豆混种时,飞机草的株高、地下生物量比及根冠比显著降低,假地豆的株高无显著变化,但生物量显著增加;飞机草的竞争参数相对产量(RY)显著小于1,竞争攻击力系数显著小于零,表明其竞争力弱于假地豆。飞机草与白饭树混种时,飞机草的根冠比也显著降低,但株高和生物量均显著增加,而白饭树的株高和生物量却显著降低;飞机草的相对产量(RY)显著大于1,竞争攻击力系数显著大于零,表明其竞争力强于白饭树。结合以上结果,本地植物假地豆可以一定程度上竞争抑制飞机草的生长,具有替代控制飞机草的潜力,而白饭树可以在清除飞机草后的入侵地辅助植被修复。实验结果为飞机草替代控制的目标物种的筛选提供了科学依据,对被飞机草入侵生境生物多样性的恢复和重建具有一定指导意义。     

Effect of herbivory on growth and biomass allocation of Brazilian peppertree (Sapindales: Anacardiaceae) seedlings in the laboratory

Brazilian peppertree (Schinus terebinthifolius Raddi), native to South America, is invading many ecosystems in south and central Florida. The defoliating tortricid moth Episimus unguiculus Clarke was selected as a potential biocontrol agent of Brazilian peppertree in Florida. The objective of this study was to examine the effect of different levels of herbivore damage on growth and biomass allocation of Brazilian peppertree seedlings in the laboratory. Three treatments were established: (1) no herbivory (control), (2) low herbivory (~4 larvae/plant), and (3) high herbivory (~12 larvae/plant). High levels of herbivory significantly reduced the number of leaflets, plant height, foliar biomass, foliar relative growth rate (RGR) and shoot: root ratio of Brazilian peppertree seedlings. Moreover, plants were not able to recover from herbivory after 2 months. The performance of Brazilian peppertree subjected to low herbivory levels did not differ from the control plants (no herbivory). The potential effectiveness of the biocontrol agent E. unguiculus to suppress this noxious weed is examined.     

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.     

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.     

Effect of organic fertiliser and Chromolaena odorata residue on the pathogenicity of Meloidogyne incognita on maize

Pot experiments laid out in a complete randomised design were conducted in the screen house of the Department of Crop Protection, University of Agriculture, Abeokuta, Ogun State, Nigeria to determine the effects of organic fertiliser and Chromolaena odorata residue at 1% w/w on the pathogenicity of Meloidogyne incognita infecting maize. M. incognita significantly reduced the plant height, number of leaves per plant, leaf area, cob weight and grain yield of maize by 6.89, 15.18, 20, 63.92 and 56.16% respectively. C. odorata residue and organic fertiliser significantly suppressed M. incognita galling, inhibited the nematode fecundity and reduced the number of eggs and juveniles on maize. A remarkable increase in plant height, number of leaves per plant, leaf area, cob weight and grain yield were observed on maize plants treated with the mixture of C. odorata and organic fertiliser despite the nematode infection. The observation from this study suggests that C. odorata in combination with organic fertiliser is a viable option for the control of M. incognita on maize.     

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.     

Invaded habitat incompatibility affects the suitability of the potential biological control agent Listronotus sordidus for Sagittaria platyphylla in South Africa

Sagittaria platyphylla (Engelmann) J.G. Smith (Alismataceae) was first recorded in South Africa in 2008 and is considered to be an emerging weed with naturalised populations occurring throughout the country. A biological control programme was initiated in Australia and surveys conducted between 2010 and 2012 yielded potential agents, including the crown feeding weevil, Listronotus sordidus Gyllenhal (Coleoptera: Curculionidae). The potential of L. sordidus as a candidate biological control agent against S. platyphylla in South Africa was examined. Although adult feeding was recorded on a number of plant species, oviposition and larval development indicated a narrow host range restricted to the Alismataceae. In South Africa, S. platyphylla populations are primarily found in inundated systems. However, laboratory studies showed that L. sordidus did not oviposit on inundated plants, potentially nullifying the impact of the insect on South African populations. It is suggested that even though L. sordidus is a damaging, specific agent, its limited impact on inundated plant populations in South Africa does not justify the inherent risk associated with the release of a biological control agent.     

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.     

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.     

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.     

Genetic variation in host susceptibility to attack by the mahogany shoot borer, Hypsipyla grandella (Zeller)

1 Attack by shoot borers (Hypsipyla grandella Zeller) is the main factor limiting the cultivation of Cedrela odorata and Swietenia macrophylla, two economically important members of the mahogany family. No viable methods of pest control are currently available. To assess for genetic variation in susceptibility to pest attack, a combined progeny/provenance test of C. odorata and a provenance test of S. macrophylla were established separately at CATIE, Turrialba, Costa Rica, and assessed intensively over an 84-week period. 2 Variation in height growth between provenances was highly significant in both species, provenance means varying by a factor of 2.7 and 1.2 in C. odorata and S. macrophylla, respectively. Cedrela odorata also displayed provenance variation in foliar phenology, as 94.7% of the trees from a ‘wet zone’ provenance remained foliated, whereas > 35% of trees from three ‘dry zone’ provenances abscised their leaves during the dry season. 3 Both species displayed significant genetic variation in susceptibility to shoot borer attack. At the peak of attack during the second year of growth, the effect of provenance was highly significant in both species, the mean number of attacks per tree varying between 0.8–2.4 and 0.6–1.3 in different provenances of C. odorata and S. macrophylla, respectively. A provenance of C. odorata from San Carlos, Costa Rica, displayed consistently lower susceptibility to pest attack, being subjected to fewer attacks during the first year and demonstrating a higher mean height to first point of damage. 4 The nitrogen, total tannin and proanthocyanidin concentration of foliage varied significantly between C. odorata provenances: nitrogen concentration was significantly lower and tannin and proanthocyanidin contents were significantly higher in trees from the San Carlos provenance, Costa Rica. This study therefore suggests that proanthocyanidins may reduce susceptibility of C. odorata to H. grandella, at least during the early period of growth. 5 These results provide support for the development of plant resistance as a strategy for managing shoot borers. In particular, there may be scope for selecting for high foliar proanthocyanidin content and the ability to tolerate attack by vigorous apical growth.     

A Plague on Your House? Some Impacts of Chromolaena odorata on Timorese Livelihoods

Following its introduction on the island of Timor, the shrub Chromolaena odorata has expanded dramatically across the landscape. A highly flammable but fire tolerant plant species, Chromolaena is an extremely successful plant coloniser of disturbed ground and open savanna woodlands. For Timorese semisubsistence agriculturalists, Chromolaena odorata represents the most recent in a long history of invasive weed species that have periodically covered their lands and challenged their ability to farm and prosper. Already its impact is being felt in a number of domains of socioeconomic life. This paper explores some of these issues and the prospects for future management and control strategies.     

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.     

Phenology of Tick weed (Cleome viscosa L.) and its interaction with okra (Abelmoschus esculentus (L.) Moench) and Melodoigyne incognita

Cleome viscosa is an emerging weed with the potential of interfering with okra and influencing pests of okra. Screen house studies were conducted on the phenology of C. viscosa, its interference with okra and its interaction with root-knot nematode-infected okra. Seedlings of C. viscosa were monitored in pots for growth, yield and dry matter accumulation for 14 weeks. C. viscosa was planted with okra at densities 0, 2, 4, 6, 8 and 10 weeds per okra plant and observed for 11 weeks. Data were collected on growth, yield and dry matter of okra. Plants were inoculated with 2,500 M. incognita eggs per pot while control plants were not inoculated. C. viscosa attained 91.7 cm height and accumulated 7.8 g/plant biomass at 14 weeks after planting. The percentage reduction in okra plant height, plant dry weight and fruit yield due to interference at lowest cleome density (2 plants/pot) was 33.7, 83.6 and 82.1%, respectively. Nematode reproductive factor was significantly lower for okra alone (4.9) compared to okra with cleome (7.6). This study shows that C. viscosa is a fast-growing weed that suppressed the performance of okra even at low density, is a good host to M. incognita and increased the population of the nematode in soil.     

Phenology of Parasetigena silvestris (Diptera: Tachinidae), gypsy moth (Lymantria dispar) (Lepidoptera: Lymantriidae) larval parasitoid and its efficiency for parasitisation

Parasetigena silvestris is a univoltine, solitary, larval endoparasitoid which lays its eggs on the surface of gypsy moth larvae. Field collection of the host larvae (2nd through 5th instar) from an artificially established gypsy moth population were made to compare stage specific parasitism between larvae without and with P. silvestris tachinid eggs. The tachinid oviposition rate detected was highest in second instar larvae, and then decreased as larvae developed toward full maturity. The opposite was true for tachinid parasitoid emergence which had no emergence from second through third host instar larvae. Fourth instar gypsy moth larvae, however, experienced significantly higher parasitism by P. silvestris in the larvae with eggs than those without the eggs. The braconid wasp Cotesia melanoscelus caused significantly higher parasitism in early instar larvae with P. silvestris eggs than in those without the eggs. The tachinid prefers to lay more eggs on parasitised larvae by the braconid even though the braconid is a superior competitor to the fly during multiparasitism. Factors influencing parasitism rates by P. silvestris such as host-parasitoid synchronisation and the multiparasitism interaction with C. melanoscelus are discussed.