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The Initiation of Biological Control Programmes Against Solanum Elaeagnifolium Cavanilles and S. Sisymbriifolium Lamarck (Solanaceae) in South Africa

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The Initiation of Biological Control Programmes Against Solanum Elaeagnifolium Cavanilles and S. Sisymbriifolium Lamarck (Solanaceae) in South Africa The initiation of biological control programmes against Solanum elaeagnifolium Cavanilles and S. sisymbriifolium Lamarck (Solanaceae) in South Africa T. Olckers1, J.H. Hoffmann2, V.C. Moran2, F.A.C. Impson2 & M.P. Hill3 1ARC - Plant Protection Research Institute, Private Bag X6006, Hilton, 3245 South Africa 2Zoology Department, University of Cape Town, Rondebosch, 7700 South Africa 3ARC - Plant Protection Research Institute, Private Bag X134, Pretoria, 0001 South Africa Two herbaceous Solanum species, Solanum elaeagnifolium Cavanilles and S. sisymbriifolium Lamarck have become problematic weeds in South Africa. A biological control programme was initiated against S. elaeagnifolium in the 1970s when a fruit-galling gelechiid moth, Frumenta nephelomicta Meyrick, was released on the weed but failed to establish. Subsequently, two chrysomelid species, Leptinotarsa texana Schaeffer and L. defecta Stål, were released on S. elaeagnifolium during 1992 and a third chrysomelid, Gratiana spadicea (Klug), was released on S. sisymbriifolium in 1994. Releases of the three beetle species were approved even though laboratory-based host-specificity studies showed that each was able to feed and develop on several native Solanum species and on a crop, S. melongena Linnaeus (eggplant). The rationale used to justify the release of these species, all of which have since become established, is discussed. Leptinotarsa texana reaches very high population densities on S. elaeagnifolium and causes considerable damage, stunting vegetative growth and fruit-production capacity. Early indications are that L. texana has the potential to contribute substantially to the biological control of S. elaeagnifolium. Conversely, L. defecta has remained localized and relatively scarce, with no obvious impact on the weed. Gratiana spadicea has become established on S. sisym- briifolium at a few release sites, but extreme winter temperatures and frequent ‘veld’ fires seem to have kept the beetle populations at low levels. After a slow start, the biological control programmes against S. elaeagnifolium and S. sisymbriifolium have made encouraging progress. Further studies are needed to determine the effect of L. texana damage on the population density of S. elaeagnifolium and to explain the limited abundance of L. defecta and G. spadicea. Key words: biological weed control, Gratiana spadicea, Leptinotarsa defecta, Leptinotarsa texana, Solanum elaeagnifolium, Solanum sisymbriifolium. The biological control campaign against weeds of S. elaeagnifolium. They noted several factors that the genus Solanum (Solanaceae) was initiated in have slowed progress with the programme the early 1970s when it became apparent that against S. elaeagnifolium. In the mid 1980s, two other control methods were largely ineffective weeds of South American origin, Solanum mauri- against species such as Solanum elaeagnifolium tianum Scopoli and S. sisymbriifolium Lamarck, Cavanilles (silverleaf nightshade, satansbos; were also targeted for biological control in South Fig. 1) (Olckers & Zimmermann 1991). This North Africa. The programme against S. mauritianum was American plant is a major weed of agriculture and reviewed by Olckers & Zimmermann (1991) and is has invaded both arable and pastoral lands in updated in this issue, while that against S. sisym- many regions of South Africa (Wassermann et al. briifolium is reviewed here for the first time. 1988; Fig. 2). Propagation is mostly vegetative and Solanum sisymbriifolium (wild tomato, sticky the ability of the weed to regenerate from small nightshade; Fig. 3), a native of warm climes in tem- fragments of root had confounded attempts at perate South America, has been present in South mechanical or herbicidal control. Africa for around 100 years, but only during the In an earlier review, Olckers & Zimmermann last decade has it been recognized as a trouble- (1991) discussed the merits of a suite of potential some invader of agricultural lands and forestry agents that might be useful for biological control of plantations (Hill et al. 1993; Hill & Hulley 1995). 56 African Entomology Memoir No. 1 (1999): 55–63 50 mm Solanum elaeagnifolium. (Drawn by A. Walters, National Botanical Institute, Pretoria.) Although the weed has a limited distribution in initiated during 1989 (Hill & Hulley 1995). At the South Africa (Fig. 2), there are several localized time, the natural enemies associated with S. mauri- dense infestations, particularly in the high-lying tianum in South America were being surveyed by regions along the eastern escarpment of Mpuma- South African entomologists and the opportunity langa. Propagation, which occurs largely through was taken to collect natural enemies on S. sisym- seeds, is enhanced by high fruit production, dis- briifolium. persal by frugivorous birds and a resilient In this paper, we review the status of the bio- seed-bank. However, the weed can also propagate control programmes against S. elaeagnifolium and vegetatively and coppices readily after mechani- S. sisymbriifolium in South Africa and describe the cal clearing which, despite offsetting fruit produc- events that permitted the releases of seemingly tion, provides inadequate control. Only one non-specific agents. These projects are the first herbicide, Garlon 4 (triclopyr), is registered for use biological control programmes against solana- against the weed. A biological control programme ceous weeds anywhere in the world and as such against S. sisymbriifolium was opportunistically are unique. Olckers et al.: Biological control of Solanum elaeagnifolium and S. sisymbriifolium 57 Distribution of Solanum elaeagnifolium (❍)and S. sisymbriifolium (●)in South Africa. (Drawn by L. Henderson, Plant Protection Research Institute, Pretoria.) EXPANDED HOST RANGES OF so far evaluated as candidate agents for S. mauri- CANDIDATE AGENTS tianum, only one species has proved host-specific The selection and introduction of potential and suitable for release (Olckers, this issue) and biocontrol agents on S. elaeagnifolium and S. sisym- both insects evaluated for S. sisymbriifolium have briifolium has been hampered by problems in displayed expanded host ranges. determining the host specificity of the agents. During host-specificity tests, the candidate Almost all of the agents tested so far have dis- agents seldom survived on plants outside the played expanded host ranges under confined genus Solanum, but some of the insect species experimental conditions and have fed on closely developed on potato (S. tuberosum Linnaeus) and related plant species that are never attacked under virtually all developed on cultivated eggplant natural conditions (Neser et al. 1990; Olckers & (S. melongena Linnaeus) and certain native conge- Zimmermann 1991; Olckers 1996). Additional neric plants, particularly in starvation tests. These complications have been the large number of results indicated that, under the confined condi- agronomic crops in the family Solanaceae, two of tions of host-specificity tests in quarantine, very which (potato and eggplant) also belong to the few solanaceous insects are likely to demonstrate genus Solanum, and the presence of at least 30 their actual host specificity. Despite the lack of indigenous species of Solanum in South Africa. evidence of attacks on solanaceous crops, particu- The dilemma this has caused was exemplified by larly eggplant, by the agents in their countries of the programme against S. elaeagnifolium where, of origin and the selection of their natural hosts eight potential agent species that were investi- during choice tests, all applications for permission gated between 1974 and 1991 (Olckers & Zimmer- to release solanaceous agents were withheld until mann 1991; Olckers 1996), only two were released. it was realized that conventional tests would The problem was exacerbated by the fact that never resolve the problem. neither species became established when released In the early 1990s, risk assessments were carried (Olckers 1995). Similarly, of the nine insect species out on two leaf-feeding chrysomelid beetles, 58 African Entomology Memoir No. 1 (1999): 55–63 50 mm Fig. 3 Solanum sisymbriifolium. (Drawn by B. Connell, National Botanical Institute, Pretoria.) Leptinotarsa texana Schaeffer and L. defecta Stål, but cause negligible damage relative to generalist which at the time were regarded as the most pests implied that imported agents were not a promising agents for biological control of significant additional risk. Furthermore, the S. elaeagnifolium. The risk to eggplant cultivations intensive pesticide regimes needed to secure the in South Africa was assessed by evaluating cultiva- crop against other pests provides a ready-made tion practices, damage inflicted by native deterrent to the introduced biocontrol agents. solanaceous insects on the crop and the nature of Despite native Solanum species not having the crop-protection procedures (Olckers & Hulley benefit of chemical protection, there was also suffi- 1994). The results provided convincing evidence cient evidence to indicate that these would not that eggplant crops were not threatened by the suffer more than incidental damage. Extensive two Leptinotarsa species (Olckers & Hulley 1994; surveys of both native and introduced Solanum Olckers & Zimmermann 1995; Olckers 1996). In species in South Africa (Olckers & Hulley 1989, particular, observations that several South African 1991, 1995; Hill et al. 1993) showed that very few of solanaceous insects feed on cultivated eggplant the native insects attack
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