Biology and host range of Ophiomyia camarae, a biological control agent for Lantana camara in South Africa

The herringbone leaf-mining fly, Ophiomyiacamarae Spencer, is a promising candidateagent for the biological control of the alieninvasive weed Lantana camara L. in SouthAfrica. During extensive host-specificity testsinvolving 39 test plant species from 12families, survival to adulthood was restrictedto L. camara, L. trifolia, and fourspecies of the closely-related genus Lippia (Verbenaceae) in no-choice tests. However, survival of the immature stages wassignificantly lower on L. trifolia andthe four Lippia species than on L.camara. In addition, O. camaraedisplayed very strong oviposition preferencefor L. camara during paired-choice tests.This narrow laboratory host range suggests thatO. camarae will not pose any risks tonon-target verbenaceous plants if released inSouth Africa. Permission to release O. camaraeinto South Africa was approved by the regulatoryauthorities in 2001.     

Distribution and abundance of the herringbone leaf-miner,Ophiomyia camarae (Diptera: Agromyzidae) on Lantana camara (Verbenaceae) in selected areas of Swaziland

The distribution and abundance of Ophiomyia camarae was determined in three ecological regions, i.e., highveld, middleveld, and lowveld of Swaziland. Within each region, sites were selected based on Lantana camara L. density, with 10 plants randomly chosen per sampling site. Ten leaves from each shrub were collected and assessed for any damage caused by the herringbone leaf-miner. Relative abundance, based on the number of shrubs damaged by the herringbone leaf-miner, was scored per site. Three lantana varieties were observed during the sampling period. There were significant differences in mean leaf damage due to leaf-miner activity between varieties (P=0.0162), with 46±4.89% leaves of the pale pink variety damaged compared to 23±5.56 and 21±3.53% leaves of the orange and pink varieties, respectively. While there were no restrictions in terms of distribution, the highest levels of O. camarae populations were observed in the middleveld (P=0.0012), suggesting that this region had the most suitable environmental conditions for this natural enemy. Overall, the abundance was significantly different between regions (P=0.006) and months (P=0.000), with April scoring the highest population levels. Based on our observations, the variation in environmental conditions has an influence on the activity of O. camarae, with the middleveld having the most suitable conditions     

Suppression of growth and reproductive capacity of the weed Lantana camara (Verbenaceae) by Ophiomyia camarae (Diptera: Agromyzidae) and Teleonemia scrupulosa (Heteroptera: Tingidae)

An 8-month study was conducted to determine the effect of herbivory by the herringbone leafminer Ophiomyia camarae Spencer (Diptera: Agromyzidae) and the lace bug Teleonemia scrupulosa Stål (Heteroptera: Tingidae) on growth and reproductive capacity of their host plant, Lantana camara L. (Verbenaceae). Ophiomyia camarae adults were released into field cages at two population densities while T. scrupulosa, a widespread and naturalised biocontrol agent in South Africa, colonised and attacked uncaged plants of similar size grown under similar field conditions. At lower initial density, O. camarae reduced stem height, stem diameter, leaf density, flower density and above-ground biomass by 5, 22, 54, 100 and 41%, respectively. At higher initial density of O. camarae, stem height, stem diameter, leaf density, flower density and above-ground biomass were substantially reduced by 19, 28, 73, 99 and 49%, respectively. Teleonemia scrupulosa also reduced stem height, stem diameter, leaf density, flower density and above-ground biomass by 37, 34, 75, 100 and 50%, respectively. The root development was less affected by both O. camarae and T. scrupulosa. Whilst higher densities of O. camarae exerted noticeable herbivore pressure, T. scrupulosa herbivory substantially hampered plant growth and reproductive capacity even at lower population levels. These findings suggest that O. camarae, released in 2001 for the control of lantana, would be effective as part of a complementary guild of biological control agents. The profound effect of herbivory by T. scrupulosa on plant growth and reproductive capacity suggests that it is making a much greater contribution to the biological control of lantana in South Africa than was previously thought.     

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.     

Tsetse flies are attracted to the invasive plant Lantana camara

In tsetse both sexes feed exclusively on the blood of vertebrates for a few minutes every 2-3 days. Tsetse flies seek cover from high temperatures to conserve energy and plants provide shelter for tsetse in all the biotopes they occupy. Recently, tsetse have taken cover in plantations and under the invasive bush Lantana camara that has invaded large areas of the tsetse fly belt of Africa. Flies from such refugia are implicated in sleeping sickness epidemics. In a wind tunnel we show that both foliage and an extract of volatiles from foliage of L. camara attract three tsetse spp. from different habitats: Glossina fuscipes fuscipes (riverine), G. brevipalpis (sylvatic) and G. pallidipes (savannah). Gas chromatography analysis of volatiles extracted from leaves and flowers of L. camara coupled to electroantennograme recordings show that 1-octen-3-ol and beta-caryophyllene are the major chemostimuli for the antennal receptor cells of the three tsetse spp. studied. A binary mixture of these products attracted these flies in the wind tunnel. The gas chromatography linked electroantennograme analysis of the L. camara extracts also show that the antennal receptor cells of the three tsetse spp. respond similarly to groups of volatiles derived from the major biosynthetic and catabolic pathways of plants, i.e. to mono- and sesquiterpenes, to lipoxidation products and to aromatics. Mixtures of these plant volatiles also attracted tsetse in the wind tunnel. These findings show that tsetse flies have conserved a strong sensitivity to volatile secondary products of plants, underlining the fundamental role of vegetation in tsetse survival.     

入侵植物马缨丹研究进展

马缨丹原产于热带美洲,现已扩散至世界热带和亚热带地区,并严重威胁入侵地本地植物的生长。马缨丹因具有观赏价值而被我国大量引进,但由于疏于管理,发展成为入侵杂草,对我国农业、畜牧业及生态环境造成严重危害。目前对马缨丹的入侵机制还没有全面系统的认识。本文从马缨丹的生物学特性、入侵机制、生态学效应、防治与应用等方面进行综述,总结其研究进展,以期为马缨丹的防治工作提供依据。研究结果显示:马缨丹成功入侵的关键因素源于其独特的生物学性征;对农林业造成的危害的原因主要为它的化感作用;马缨丹的防治方法主要包括物理、化学和生物防治;马缨丹在防虫除虫、污染修复、药用价值等方面有所应用。此外,分析当前研究中的不足后发现,今后应加强马缨丹在自然环境中的适应机制、应用潜力、防治方法等的研究。     

Geographical range and impact of five biocontrol agents established on Lantana camara in South Africa

A survey was conducted todetermine the present status of five biocontrolagents established on Lantana camara L.(Verbenaceae) in South Africa. Three ofthe five agents, Calycomyza lantanae(Frick), Ophiomyia lantanae Froggatt andTeleonemia scrupulosa Stål areestablished throughout the range of L.camara. The two hispine beetles, Octotomascabripennis Guérin-Ménevilleand Uroplata girardi Pic are restrictedto the warm, subtropical regions in the easternrange of the weed, and are unable to cope withthe plants becoming seasonally leafless in dryand temperate areas. The two beetles and T. scrupulosa are rated as the most damagingagents on L. camara. The impact of O. lantanae is uncertain, and due to lowpopulations, the impact of C. lantanae isnegligible. Insect populations typicallyaccumulate to maximise their impact on plantsby midsummer, giving plants the opportunity tocompensate for the cumulative agent damageaccrued at the end of the previous growingseason. A number of parasitoids were rearedfrom U. girardi, O. lantanae andC. lantanae, but only those adapted toC. lantanae are expected to significantlyreduce its field density. The agents feed anddevelop on a wide range of lantana varieties,and where two or more varieties co-exist,insects occurred in equal abundance on thedifferent varieties. The importance of varietalpreferences appears to have beenover-estimated, but needs consideration whennew candidates are evaluated. The impact ofthese biocontrol agents is insufficient toreduce L. camara to a manageablesituation, and additional candidates arenecessary to control this weed in South Africa.     

Contrasting strategies to cope with drought by invasive and endemic species of Lantana in Galapagos

This study compares how Lantana camara, an invasive species, and L. peduncularis, an autochthonous one, cope with drought in Galapagos. Soil surface temperature was the abiotic environmental parameter that best explained variations in photosynthetic stress. Higher soil surface temperatures were recorded in the lowlands and in rain-shadow areas, which were also the driest areas. L. peduncularis, with a shallow root system, behaved as a drought-tolerant species, showing lower relative growth rates, which decreased with leaf water content and higher photosynthetic stress levels in the lowlands and in a northwest rain-shadow area in comparison with higher and wetter locations. Its basal and maximal fluorescences decreased at lower altitudes, reflecting the recorded drops in chlorophyll concentration. In contrast, L. camara with a deep root system behaved as a drought-avoiding species, showing leaf and relative water contents higher than 55% and avoiding permanent damage to its photosynthetic apparatus even in the driest area where it showed very low chlorophyll content. Its relative growth rate decreased more in dry areas in comparison to wetter zones than did that of L. peduncularis, even though it had greater water content. Furthermore, L. camara showed higher water contents, growth rate, and lower photosynthetic stress levels than L. peduncularis in the arid lowlands. Thus, L. peduncularis maintained lower maximum quantum efficiency of photosystem II photochemistry (F _v/F _m) than L. camara even at sunrise, due to higher basal fluorescence values with similar maximal fluorescence, which indicated permanent damage to PSII reaction centres. Our results help to explain the success and limitations of L. camara in the invasion of arid and sub-arid environments.     

Status, invasiveness and environmental threats of three tropical American invasive weeds (Parthenium hysterophorus L., Ageratum conyzoides L., Lantana camara L.) in India

Invasive weeds have threatened the integrity of ecosystems throughout the world. They affect not only the species diversity of native areas but also their biological integrity. In India, a number of invasive exotic weeds have been reported but some viz. Parthenium hysterophorus, Lantana camara and Ageratum conyzoides, especially those from tropical America are troublesome and have caused adverse ecological, economic and social impact. These weeds can be seen growing in different landscapes but are luxuriantly localized in unattended forests and cultivated areas. Parthenium hysterophorus (Asteraceae, commonly known as congress grass) is perhaps the most troublesome and noxious weed of urban and rural India. Besides rapidly colonizing areas replacing the native vegetation, it is also known to cause a number of human health problems such as skin allergy, rhinitis and irritation to eyes of the residents in the vicinity. Likewise, it causes fodder scarcity in addition to being unpalatable and toxic to livestock. Lantana camara (Verbenaceae), another serious tropical American pest, has encroached upon large areas of land, especially the forests where it has virtually replaced the forest floor vegetation and reduced tree growth. Also because of its bushy and spreading type of growth it obstructs forest operations. The third weed, Ageratum conyzoides (Asteraceae, Billy goat weed) has invaded agricultural fields. It interferes with crops and causes yield reductions of major staple crops of India. When it invades rangeland areas, it out competes native grasses causing scarcity of fodder. These weeds have similar growth strategies such as fast growth rates, short life-cycles, greater reproductive potential, high competitive abilities and allelopathy that make them successful invaders of native habitats. Mechanical, chemical, biological and cultural control tactics have failed individually, though integrated approaches combining all these approaches along with community participation and proper land management have been relatively successful. This paper presents various aspects of biology, ecology, hazards and control measures of these weeds.     

Fungal pathogens of Euphorbia heterophylla and E. hirta in Brazil and their potential as weed biocontrol agents

A two-year survey of the fungi associated with two important congeneric pantropical weeds, Euphorbia heterophylla and E. hirta, was conducted in part of their native range in southern Brazil. Sampling was concentrated mainly in Rio de Janeiro State and ten species were identified as pathogens of these weeds. Two taxa, Botrytis ricini and Uromyces euphorbiae, were common to both weed hosts. Alternaria euphorbiicola, Bipolaris euphorbiae, Melampsora sp., Oidium sp. and Sphaceloma poinsettiae were recorded only from E. heterophylla, whereas Colletotrichum gloeosporioides, Sphaceloma sp. and Sphaerotheca fuliginea were restricted to E. hirta. Botrytis ricini and Colletotrichum gloeosporioides are new records for E. hirta, and Alternaria euphorbiicola and Sphaerotheca fuliginea are new host records for Brazil. Bipolaris euphorbiae, previously identified as Helminthosporium sp., is considered to be the correct name for the causal agent of a major disease of E. heterophyllum in Brazil. The potential of these pathogens as biocontrol agents is discussed and the mycobiota associated with both these weeds worldwide is reviewed.This revised version was published online in October 2005 with corrections to the Cover Date.     

Biological control of weeds by means of plant pathogens: Significance for integrated weed management in modern agro-ecology

Biological control of weeds by using plant pathogens has gainedacceptance as a practical, safe, environmentally beneficial, weedmanagement method applicable to agro-ecosystems. The interest in thisweed control approach from public and private groups, and support forresearch and developmental effort, are on the upswing. This increasinginterest is stimulated largely due to major economic, social, andenvironmental forces that are directing our choices in crop productionpractices. Some of these changes are market-driven while others aresocial and ecological in nature. These changes are in turn influencingthe choices in weed control methods. In this regard, biocontrol withplant pathogens has been proven a feasible, albeit minor, component ofmodern integrated weed-management systems. This environmentallybeneficial method should be promoted and exploited further to meet thecurrent and future challenges in weed management in agro-ecosystems.     

A pilot project testing the effectiveness of three weed control methods on the removal of Lantana camara in Forty Mile Scrub National Park,Queensland, Australia

Lantana (Lantana camara) poses a serious threat to the biodiversity of the dry rainforest vegetation at Forty Mile Scrub National Park, Queensland, Australia, by outcompeting native species and increasing vulnerability to fire. This pilot study tests the effectiveness of three weed control methods (hand pulling, a glyphosphate‐based foliar spray herbicide and a picloram‐ and triclopyr‐based basal bark herbicide) in removing lantana and their success in reducing lantana fuel loads. The foliar spray herbicide was the most effective in killing lantana, while manual pulling resulted in the largest decrease in fuel height. We suggest that foliar spraying will be most efficient for combating large infestations of lantana, while hand pulling techniques are recommended for creating firebreaks or when minimizing damage to native species is paramount.     

Unsuitability of Charidotis pygmaea for biocontrol of Lantana camara in Africa

The host range of the tortoise beetle,Charidotis pygmaea Klug (Coleoptera:Chrysomelidae), was studied under quarantinelaboratory conditions to evaluate the insect'ssuitability for release as a biological controlagent for the noxious weed, Lantanacamara L. (Verbenaceae) in South Africa.Culturing on the target plant, L. camara,proved problematic with high larvalmortalities. Host-specificity studies showedthat four species in the genus Lantana,and two species in the genus Lippiawere acceptable as host plants. Duringlarval development trials, the insect performedbetter on the indigenous Lantana rugosaThunb. (Verbenaceae) and the introduced,commercially used L. montevidensis(Spreng.) Briq. (Verbenaceae), than on any ofthe weedy South African L. camaravarieties tested. Adult multi-choice trialsindicated that the beetle preferred to ovipositon L. rugosa and L. montevidensis.It is therefore recommended that C.pygmaea not be released against L.camara in Africa.     

First record of Cercospora pistiae causing leaf spot of water lettuce ( Pistia stratioites ) in Brazil,with particular reference to weed biocontrol

Pistia stratioites , the water lettuce, is an important pantropical aquatic weed. It has been the target of successful classical biocontrol programmes utilizing an insect natural enemy collected in Brazil and introduced into Australia, Papua New Guinea and South Africa. Little attention was given to the fungal pathogens associated with this weed as potential biocontrol agents. An 11-month field survey in the state of Rio de Janeiro involving repeated visits to 12 sites, and a 2-month survey in the Amazonas Basin (Amazonas state) in Brazil, yielded only one pathogenic fungus on P. stratioites. This was identified as Cercospora pistiae. No previous record of this species has been made in Brazil nor in the New World. The fungus is described and its taxonomy and biocontrol potential discussed herein. C. pistiae is a damaging pathogen of water lettuce. Its distribution was restricted to two interconnected sites in Rio de Janeiro. A literature and herbaria survey yielded few fungi associated to water lettuce (8 species), probably reflecting a lack of adequate collection in the centre of origin of the plant. This revised version was published online in August 2006 with corrections to the Cover Date.     

Biological control of <Emphasis Type="Italic">Lantana camara</Emphasis> in South Africa: targeting a different niche with a root-feeding agent, <Emphasis Type="Italic">Longitarsus</Emphasis> sp.

The root-feeding flea beetle, Longitarsus sp. (Coleoptera: Chrysomelidae: Alticinae), was studied as a potential biological control agent for Lantana camara L. (Verbenaceae) in South Africa. Host range tests were carried out on 52 plant species in 11 families. Although 11 plant species, all in the family Verbenaceae, supported complete development of Longitarsus sp. during no-choice tests, the beetles showed very strong preferences for L. camara during paired-choice and multi-choice tests. The results confirm that the beetles have a narrow host range, and that under natural conditions they are highly unlikely to utilise plants other than L. camara. In the unlikely event that some of the Lippia spp. are attacked in the field, they are not expected to sustain populations of the flea beetle over time. Attributes that should enhance the biocontrol potential of Longitarsus sp. include: the adults are long-lived and highly mobile; and, the larvae cause extensive direct damage to the roots of L. camara, which could in turn expose the plants to soil-born pathogens. All indications are that Longitarsus sp. could make a substantial contribution to the biological control of L. camara in many countries around the world because the beetles pose no threat to non-target plant species and they damage a part of the plant (i.e. roots) not yet affected by any other agent species.     

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.     

Ontogenesis of internal secretory cells in leaves of Lantana camara (Verbenaceae)

Internal secretory cells may be morphologically indistinguishable from their neighbours except for the presence of secreted material, or they may differ to such an extent that could be classified as secretory idioblasts. Several authors have reported the presence of glandular trichomes secreting essential oil in Verbenaceae, including Lantana . However, none have reported internal secretory cells. Anatomical and histochemical methods applied to Lantana camara leaves revealed the occurrence of internal secretory cells whose ontogenesis and chemical nature are described in this paper. According to leaf developmental analysis, L. camara secretory cells originated from the ground meristem, started to differentiate in the third node leaves, and were actively secreting in the fourth node leaves. The content of the secretory cells was of a lipidic nature, and a terpenoid essence of their secretion was also identified. Based upon differences in size and shape from neighbouring cells and on detection of nonvolatile terpenes, they were confirmed as true internal secretory idioblasts.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 427–431.     

Antitermitic effect of the Lantana camara plant on subterranean termites (Isoptera: Rhinotermitidae)

Abstract The insecticidal effects of Lantana camara L. (flowers, leaves, stems and roots) and the soil where lantana had been growing, on foraging activity and survival of the subterranean termites Coptotermes formosanus and Reticulitermes flavipes were examined in a 3-week experiment. The soil in which lantana had been growing had no effect on termite tunneling and survival. Incorporation of chipped fresh lantana leaves and stems into soil had no effect on mortality but caused significant reduction in tunneling. The 5-cm wide barrier of soil with lantana tissue incorporated effectively repelled groups of both species from penetrating the barrier and thus prevented infestation of a piece of wood on the other side of the barrier. C. formosanus was more sensitive in avoiding the barrier than R. flavipes. Leaves, stems and flowers were more repellent than roots. These results provide preliminary evidence that fresh-cut lantana leaves, stems and flowers may have use as additives to garden mulches against termites.     

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16′ S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.