David O. Simelane

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.

Influence of soil texture, moisture, and surface cracks on the performance of a root-feeding flea beetle, Longitarsus bethae (Coleoptera: Chrysomelidae), a biological control agent for Lantana camara (Verbenaceae).

Abstract Laboratory studies were conducted to determine the influence of soil texture, moisture and surface cracks on adult preference and survival of the root-feeding flea beetle, Longitarsus bethae Savini and Escalona (Coleoptera: Chrysomelidae), a natural enemy of the weed, Lantana camara L. (Verbenaceae). Adult feeding, oviposition preference, and survival of the immature stages of L. bethae were examined at four soil textures (clayey, silty loam, sandy loam, and sandy soil), three soil moisture levels (low, moderate, and high), and two soil surface conditions (with or without surface cracks). Both soil texture and moisture had no influence on leaf feeding and colonization by adult L. bethae. Soil texture had a significant influence on oviposition, with adults preferring to lay on clayey and sandy soils to silty or sandy loam soils. However, survival to adulthood was significantly higher in clayey soils than in other soil textures. There was a tendency for females to deposit more eggs at greater depth in both clayey and sandy soils than in other soil textures. Although oviposition preference and depth of oviposition were not influenced by soil moisture, survival in moderately moist soils was significantly higher than in other moisture levels. Development of immature stages in high soil moisture levels was significantly slower than in other soil moisture levels. There were no variations in the body size of beetles that emerged from different soil textures and moisture levels. Females laid almost three times more eggs on cracked than on noncracked soils. It is predicted that clayey and moderately moist soils will favor the survival of L. bethae, and under these conditions, damage to the roots is likely to be high. This information will aid in the selection of suitable release sites where L. bethae would be most likely to become established.

Suitability of some Lantana camara varieties as host plants for the root-feeding flea beetle, Longitarsus bethae

Abstract Compatibility between the root-feeding agent Longitarsus bethae Savini & Escalona and the varieties of the target weed Lantana camara L. could influence the chances of successful establishment and effectiveness of this agent as a biocontrol agent of lantana in South Africa. Laboratory experiments were conducted to determine the influence of major South African lantana varieties on feeding, colonization, oviposition preference and larval survival of L. bethae. Feeding preference and the number of adults per plant varied among the varieties. The most damaged and colonized varieties were 150 Orange, 009 Light Pink and 113 Dark Pink. Variety 018 Dark Pink was the least damaged, while 021 Total Pink and 029 White Pink were the least colonized. Females laid the highest number of eggs on variety 150 Orange. The lowest number of eggs was recorded on Australian Richmond Pink and the closely related species Lippia wilmsii. Larval survival differed significantly among the varieties. The highest survival was recorded on 150 Orange. Duration of development also varied significantly among the varieties, with the slowest rate of development recorded on variety 029 Light Pink and the quickest on variety 015 White Yellow. Although variations in female body size were less significant among varieties, males emerging from variety 150 Orange were slightly smaller than those emerged from other varieties. There was a significant tendency for females to select varieties that enhanced the survival of their offspring, which met the predictions of the preference-performance hypothesis. The data suggest that the chances of establishment and successful control of the weed are expected to be much greater on those varieties that were preferred for oviposition and provided better larval performance.

Interaction between Uroplata girardi (Coleoptera: Chrysomelidae) and Ophiomyia camarae (Diptera: Agromyzidae) on a shared host Lantana camara (Verbenaceae)

ABSTRACT Multiple releases of insect agents intended to target a single plant pest species could result in competitive interactions that in turn might affect the community structure of the phytophagous insects. Two leaf-feeding biological control agents, Uroplata girardi Pic (Coleoptera: Chrysomelidae) and Ophiomyia camarae Spencer (Dipetera: Agromyzidae), were released against the weed Lantana camara L. (Verbenaceae) in South Africa in the 1970s and 2001, respectively. Since the population explosion of O. camarae in 2005, a decline of U. girardi populations had been observed in KwaZulu-Natal (KZN) humid coast, leading to speculation that negative interaction may be operating between the agents. The study therefore was conducted to determine the competitive effect of O. camarae on U. girardi. The study showed that 76% of O. camarae larval mines were formed on uninfested (clean) compared with only 24% formed on U. girardi-infested leaves, suggesting that the fly chose to lay more eggs on clean leaves. Almost the same number of U. girardi larval mines was formed on both O. camarae-infested and clean leaves, indicating that U. girardi females in this case oviposited indiscriminately on the two types of leaves. The survival of U. girardi was 53.8% when reared on clean leaves compared with only 14.6% survival on O. camarae-infested leaves. At the end of the sampling period, densities of U. girardi was over two times higher in single-species than in combined-species treatment. Releasing both agents together did not significantly affect O. camarae densities during the sampling period. In the field, O. camarae densities increased rapidly from spring to autumn, whereas those of U. girardi remained consistently low during the same period. The bias toward oviposition on clean leaves in O. camarae enables its larvae to avoid unfavorable encounters with U. girardi larvae, thus enhancing its development and survival. The apparent inability of U. girardi to distinguish between suitable and unsuitable leaves for oviposition could compromise the fitness of this beetle, and this could explain the suppression of U. girardi populations during summer when O. camarae populations begin to increase rapidly. This study provides evidence for an asymmetric interaction between two introduced agents, and therefore highlights the importance of conducting interaction studies on agents with extensive niche overlap before their release into the environment.

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

Abstract 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.

Field and laboratory studies to determine the suitability of Cissoanthonomus tuberculipennis (Coleoptera: Curculionidae) for release against Cardiospermum grandiflorum (Sapindaceae) in South Africa

Cardiospermum grandiflorum is an invasive creeper that was targeted for biological control in South Africa in 2003. To determine ecological host range of its natural enemies, surveys were conducted on C. grandiflorum and 11 other Sapindaceae at 40 sites in the weeds native range (Argentina). These surveys indicated that the seed-feeding weevil Cissoanthonomus tuberculipennis was restricted to C. grandiflorum, and that it was among the common natural enemies, occurring at most sites where C. grandiflorum was recorded. Open-field tests were conducted under natural conditions in Argentina to determine the host preference of C. tuberculipennis and other natural enemies of C. grandiflorum among three Cardiospermum species. These tests revealed that C. tuberculipennis and the bug Gargaphia sp. were restricted to C. grandiflorum though the latter subsequently developed on non-target Cardiospermum species in the laboratory. C. tuberculipennis was found to be highly damaging, destroying up to 44% of the seeds per plant in Argentina. In all the host-specificity tests, including no-choice, paired-choice and multi-choice tests, C. tuberculipennis only fed and developed on C. grandiflorum. Failure of C. tuberculipennis to feed and develop on all congeners of C. grandiflorum shows that the weevil is highly host-specific to the target weed. Results of host-specificity tests, open field tests and long-term monitoring of C. tuberculipennis populations demonstrate that the weevil poses no threat to non-target plant species, and therefore safe for release against C. grandiflorum in South Africa. Permission to release C. tuberculipennis in South Africa has been granted by the relevant regulatory authorities.

Host-plant variety and not climate determines the establishment and performance of Aceria lantanae (Eriophyidae), a biological control agent of Lantana camara in South Africa

The flower-galling mite Aceria lantanae (Cook) (Trombidiformes: Eriophyidae) was released for the biological control of Lantana camara L. (Verbenaceae) in South Africa in 2007, but has displayed variable and patchy establishment throughout the weed’s range. Surveys were undertaken in 2013–2014, both seasonally and during the mite’s peak infestation periods, to determine the influence of climatic factors on its performance. Although there were seasonal differences in the percentages of mite-infested inflorescences, these did not differ significantly between altitudinal zones. There were also no significant relationships between the percentages of mite-infested inflorescences and either of annual rainfall, temperature or relative humidity. A field inoculation trial revealed significant differences between 10 common South African L. camara varieties in their susceptibility to A. lantanae. Only three varieties displayed appreciable susceptibility (50–61% of inflorescences infested), whereas six displayed only slight to moderate susceptibility (8–21%) and one displayed a lack of susceptibility (no infestation). These data support the contention that differential varietal susceptibility and not climate is responsible for the variable performance of A. lantanae on L. camara in South Africa. Complementing the current biotype of A. lantanae, originally sourced from Florida (USA), with other biotypes from different L. camara genotypes in Central and South America could increase the mite’s impact on the weed.

Occurrence of different Lantana camara varieties across four South African provinces and their susceptibility to a biotype of the gall-forming mite Aceria lantanae

ABSTRACT A biotype of the flower-galling mite, Aceria lantanae (Cook) (Trombidiformes: Eriophyidae) collected in Florida (U.S.A.) was released in South Africa in 2007 against Lantana camara L. (Verbenaceae) but has displayed patchy establishment. The occurrence of different L. camara varieties and their susceptibility to A. lantanae were assessed across four provinces with dense infestations. Surveys were undertaken at 113 sites during the mite’s peak infestation period (April–May) in 2013–2015. The occurrence of 13 recorded L. camara varieties differed substantially across and within these provinces. Overall, five varieties accounted for 7–45% of the sampled plants at 9–51% of the surveyed sites. The remaining eight varieties accounted for <1–4% of the plants at 2–9% of the sites. The establishment and impact of A. lantanae differed significantly between L. camara varieties. The mite established best on three varieties (163 LP, 021 WP and 015 OR), with 60–90% of plants infested. Reduced establishments were observed on seven varieties, with 18–50% of plants infested, while no establishment was recorded on three varieties. Where established, A. lantanae inflicted considerable levels of damage (i.e. 51–75% of buds infested) on the most widespread and abundant L. camara variety (163 LP) and on one less common variety (021 WP). Two uncommon varieties (015 OR, 021 P) suffered moderate levels of damage with the remainder suffering only trivial levels. The mite’s impact in South Africa could be improved by complementing the established biotype with others from Central and South America that are better matched with the poorly attacked L. camara varieties.

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

ABSTRACT 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.