Terence Olckers

Assessing the risks of releasing a sap-sucking lace bug, Gargaphia decoris, against the invasive tree Solanum mauritianum in New Zealand

The South American tree Solanum mauritianum Scopoli (Solanaceae), a major environmental weed in South Africa and New Zealand, has been targeted for biological control, with releases of agents restricted to South Africa. The leaf-sucking lace bug, Gargaphia decoris Drake (Tingidae), so far the only agent released, has become established in South Africa with recent reports of severe damage at a few field sites. To evaluate the insect’s suitability for release in New Zealand, host-specificity testing was carried out in South Africa in laboratory and open-field trials, with selected cultivated and native species of Solanum from New Zealand. No-choice tests confirmed the results of earlier trials that none of the three native New Zealand Solanum species are acceptable as hosts. Although the cultivated Solanum muricatum Aiton and S. quitoense Lam. also proved unacceptable as hosts, some cultivars of S. melongena L. (eggplant) supported feeding, development and oviposition in the no-choice tests. Although eggplant was routinely accepted under laboratory no-choice conditions in this and previous studies, observations in the native and introduced range of G. decoris, open-field trials and risk assessment based on multiple measures of insect performance indicate that the insect has a host range restricted to S. mauritianum. These results strongly support the proposed release of G. decoris in New Zealand because risks to non-target native and cultivated Solanum species appear to be negligible. An application for permission to release G. decoris in New Zealand will be submitted to the regulatory authority.

Should the flower bud weevil Anthonomus santacruzi (Coleoptera: Curculionidae) be considered for release against the invasive tree Solanum mauritianum (Solanaceae) in New Zealand?

Abstract The invasive tree Solanum mauritianum Scop. has been targeted for biological control in South Africa and New Zealand, by deploying insect agents that could constrain its excessive reproductive output. The flower-feeding weevil Anthonomus santacruzi (Curculionidae) was approved for release in South Africa in 2007 but following the loss of the original culture in quarantine, new stocks were introduced from Argentina in 2008–2009. This study was initiated to confirm that the host range of the new culture was the same as that of the previous one, but also to assess the risks associated with the weevils release in New Zealand. Different testing procedures, including no-choice tests and multi-choice tests in different arenas, produced inconsistent and ambiguous results. During no-choice tests, oviposition and larval development to adulthood occurred on three non-target species including two native South African and one native New Zealand Solanum species. However, subsequent multi-choice tests and a risk assessment suggested that the risks of anything more than collateral damage to non-target Solanum species are low. Overall, these data do not deviate substantially from the results of the original quarantine tests which facilitated the release of A. santacruzi in South Africa in 2009. Although we argue that none of the New Zealand native and cultivated species are at risk, stronger evidence from open-field trials and chemical ecology studies may be required to convince the regulatory authorities that A. santacruzi is suitable for release in New Zealand.

Genetic diversity in Pareuchaetes insulata and its implications for biological control of Chromolaena odorata

AbstractChromolaena odorata (L.) King & H.E. Robins. (Asteraceae) threatens several economic and environmental activities in the eastern subtropical regions of South Africa and is a target for biological control. Three populations of Pareuchaetes insulata (Walker) (Lepidoptera: Erebidae) originating from Florida (USA), Cuba and Jamaica were released at 30 separate sites in KwaZulu-Natal, South Africa. Subsequent overlapping of two of these populations in the field and the likelihood of interbreeding posed a potential risk of establishment failure. To explore the genetic diversity of the different P. insulata populations and test for the existence of pre- and post-zygotic reproductive isolation between them, molecular analyses and cross-mating experiments were conducted. Mitochondrial DNA sequences revealed significant genetic similarity between them. Cross-mating trials between Floridian and Jamaican populations of P. insulata revealed no significant pre- and post-zygotic isolation, with no demonstrable differences in measured parameters between the ‘parental’ populations, the F1 ‘hybrids’ and the backcrosses. These results suggest that P. insulata populations released in South Africa are part of the same ‘parental’ population. Genetic incompatibility is, therefore, relatively unlikely to be the cause of any failure in establishment of P. insulata at any of the release sites.

Does height of exposure in the canopy influence egg mortality in Acanthoscelides macrophthalmus (Coleoptera: Chrysomelidae), a biological control agent of Leucaena leucocephala in South Africa?

Abstract Introduced into several countries worldwide for agroforestry, Leucaena leucocephala (Lam.) de Wit is a typical ‘conflict species’ that invades several riparian, forested and disturbed habitats in South Africa. The seed beetle Acanthoscelides macrophthalmus (Schaeffer) (Chrysomelidae: Bruchinae) was subsequently deployed as a seed-reducing agent to offset its excessive seed output. However, exploratory egg-exposure trials, in which native Acacia s.l. trees were used as ‘surrogate host plants’, suggested that high egg mortality is reducing the beetles impact. The aims of this study were to (1) verify this suspicion by comparing the mortality of eggs exposed in L. leucocephala and Acacia nilotica (L.) Willd ex Del. stands and (2) determine whether height of exposure in the canopy (i.e. oviposition site selection) influences the different mortality factors. Although lower than in the exploratory trials, egg mortality was considerable, ranging from 30 to 50% in most instances. While unaffected by height of exposure, the overall egg mortality (i.e. all factors) was significantly lower in seeds exposed on L. leucocephala. Mortality was not only mostly attributed to parasitism (51% of mortality), but also predation (30%) and unknown factors (19%). While unexplained mortality and predation were not influenced by tree species or height of exposure, parasitism increased significantly with height above ground and was significantly lower in seeds exposed on L. leucocephala. Egg mortality is thus one of several factors that appears to be limiting the efficacy of A. macrophthalmus in South Africa.

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

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

Climate constrains the establishment and proliferation of Anthonomus santacruzi, a flower-feeding biological control agent of the invasive weed Solanum mauritianum in South Africa

ABSTRACT The flowerbud-feeding weevil Anthonomus santacruzi Hustache (Coleoptera: Curculionidae) was released in South Africa in 2008 for the biological control of the invasive tree Solanum mauritianum Scopoli (Solanaceae). The weevil was widely deployed throughout KwaZulu-Natal province, which supports large S. mauritianum infestations, and has become well established in its warmer coastal regions. The aim of this study was to provide field evidence that climate is constraining the weevil’s distribution in South Africa. Solanum mauritianum populations were sampled at 23 sites across an altitudinal gradient in KwaZulu-Natal to determine A. santacruzi densities in relation to food availability and climatic variables. Despite significantly higher amounts of floral material on S. mauritianum at the higher altitude inland sites, A. santacruzi numbers were significantly higher at the lower altitude coastal sites. There was thus a significant negative relationship between A. santacruzi numbers and altitude and significant positive relationships between A. santacruzi numbers and both temperature and humidity. Neither rainfall nor food availability influenced A. santacruzi numbers, although lower amounts of floral material at the coastal sites may well have been caused by higher weevil densities at these sites. Anthonomus santacruzi was absent at only three sites, all at higher altitudes, further demonstrating that conditions in coastal or low-altitude regions are favourable for establishment and population proliferation. Future release efforts in KwaZulu-Natal, but also in other South African provinces, should thus be focused on coastal regions and inland regions that are below 1000 m above sea level.

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.

A review of Solanum mauritianum biocontrol: prospects, promise and problems: a way forward for South Africa and globally

The invasive tree Solanum mauritianum Scopoli remains one of the world’s most widespread environmental weeds. Despite biocontrol providing one of the few viable long-term solutions to tackling S. mauritianum invasions globally, only South Africa and, more recently, New Zealand, have programmes in place. Ongoing biocontrol efforts against S. mauritianum are reviewed here with particular reference to South Africa. The South African programme has suffered a troubled history, with considerable research efforts culminating in the eventual release and establishment of only two insect agents, Gargaphia decoris Drake and Anthonomus santacruzi Hustache. The difficulties experienced have hindered research into new agents, causing apprehension in using biocontrol internationally. However, recent studies have demonstrated that biocontrol may be deserving of renewed investment, particularly within an integrated management context. In this review, we advocate for the revival of the S. mauritianum biocontrol programme in South Africa, and discuss possible avenues for future research internationally.

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.

Thermal tolerance of Liothrips tractabilis, a biological control agent of Campuloclinium macrocephalum recently established in South Africa

The South American herb Campuloclinium macrocephalum (Less.) DC. (Asteraceae; pompom weed), which invades grasslands, savannas, and wetlands in South Africa, is a recent target for biological control. A foliage‐deforming thrips, Liothrips tractabilis Mound & Pereyra (Thysanoptera: Phlaeothripidae), was released in South Africa in 2013, following confirmation of its host specificity and damage potential in quarantine. However, weed biocontrol agents sometimes fail to establish or have the desired impact because they are poorly suited to the climate of the new range. Thermal tolerance trials, within the range of 15–32.5 °C, were conducted prior to the thrips’ release to determine its developmental threshold and to generate a degree‐day model to highlight areas best suited for its establishment in South Africa. These laboratory data were validated by outdoor trials. The thrips completed development from egg to adulthood at all except the lowest and highest laboratory test temperatures, with developmental duration decreasing with increasing temperature. The lower developmental threshold was estimated at 9.6 °C with 546.9 degree days required to complete development. Within the regions invaded by C. macrocephalum in South Africa, the thrips was predicted to complete 3–9 annual generations. Developmental times in the outdoor trials were largely consistent with those in the laboratory. Liothrips tractabilis is largely compatible with the broader South African climate and seems likely to establish and proliferate on C. macrocephalum populations throughout the country. These predictions have partially been verified in that L. tractabilis has become established at several sites in South Africa, many of these in colder, high‐altitude areas which were predicted to support fewer annual generations.