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Lantana Camara (Verbenaceae) in South Africa

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Lantana Camara (Verbenaceae) in South Africa Past and present initiatives on the biological control of Lantana camara (Verbenaceae) in South Africa J-R. Baars & S. Neser ARC - Plant Protection Research Institute, Private Bag X134, Pretoria, 0001 South Africa Lantana camara, a highly invasive weed in many countries, has been targeted for biological control in South Africa since the early 1960s. An earlier review in 1991 indicated that, despite the establishment of several natural enemy species, the programme has largely been unsuccessful. In this paper we review initiatives undertaken during the 1990s and discuss (i) the status of the natural enemies established on the weed, (ii) factors that have limited the impact of these agents, (iii) the potential of eleven new biocontrol candidates currently under evaluation for re- lease and (iv) the problem of expanded host ranges of imported natural enemies under labora- tory conditions. Ultimately, the success of the programme will depend on the establishment of a suite of natural enemies, attacking several parts of the weed, which are able to cope with the ex- treme variability and wide distribution of L. camara in South Africa. Despite the problems associ- ated with the programme, L. camara remains a candidate for biological control in South Africa. Key words: Lantana camara, varieties, biological weed control, natural enemies, host-specificity testing, Lippia. Lantana camara sensu lato (Verbenaceae; Fig. 1), a compounds, notably the pentacyclic triterpenes floriferous, prickly, thicket-forming shrub, which (Kellerman et al. 1996), lantadene A and B (Morton is commonly known as lantana, originates from 1994), which if consumed can cause photosensiti- tropical and subtropical South and Central Amer- zation, liver and kidney damage, paralysis of the ica (Stirton 1977). As a popular ornamental plant, gall bladder, intestinal haemorrhage and death several varieties (cultivars) of lantana have been within 1–4 days in cattle, sheep and horses. Live- widely distributed throughout the tropics, sub- stock previously exposed to lantana are less likely tropics and warm temperate regions of the world. to suffer from the acute symptoms caused by Following its worldwide introduction, lantana has ingestion, while those with no previous exposure become naturalized in some 50 countries and is are likely to be severely affected. The expected rated as one of the world’s worst weeds (Holm annual impact of cattle mortalities from lantana et al. 1977). poisoning in South Africa was estimated to be in Lantana is an aggressive, vigorously growing excess of R1.7 million (Kellerman et al. 1996). weed that tolerates a wide variety of environmen- Lantana is an extremely variable entity that pres- tal conditions, but thrives better in humid than in ents a complex taxonomic problem (Munir 1996). dry regions. In South Africa it is presently natural- Its conspicuous variability in general morphology ized in the warm, moist subtropical and temperate (Howard 1970; Smith & Smith 1982; Cilliers 1987a; regions (Oosthuizen 1964; Stirton 1977) of the Munir 1996), physiology and genetic composition Northern Province, Gauteng, Mpumalanga and (Spies & Stirton 1982a,b,c; Spies 1984; Spies & Du KwaZulu-Natal, as well as the southern coastal re- Plessis 1987), has led to the recognition of hun- gions of the Eastern and Western Cape Provinces dreds of cultivars or varieties, all of which are clas- (Fig. 2). Lantana invades river-banks, mountain sified as L. camara. The weed is a polyploid slopes and valleys, pastures and commercial complex, presumably derived from the deliberate forests where it forms impenetrable stands that hybridization of species in the genus Lantana obstruct access and utilization. Through allelo- (Stirton 1977) and, subsequent to naturalization in pathic suppression of indigenous plant species, South Africa, hybridization was shown to con- lantana invasions also interrupt regeneration tinue in the field (Spies & Stirton 1982a,b,c). Spies processes (Gentle & Duggin 1997a) and reduce the (1984) argued that L. camara is in an active phase of biodiversity of natural ecosystems. evolution, with intermediates in a transitional Lantana leaves, stems and fruits contain toxic stage of speciation. Although the species is consid- 22 African Entomology Memoir No. 1 (1999): 21–33 50 mm Lantana camara. (Drawn by R. Weber, National Botanical Institute, Pretoria.) ered to be unstable, certain varieties dominate in strategy (Gentle & Duggin 1997b). However, the localized areas and remain relatively stable in use of fires is not always a suitable option as infes- space and time. Following ecological disturbances, tations are often close to, or in, indigenous forests, lantana stands may recolonize as a mixture of vari- grazing lands and plantations. The difficulties and eties, but soon revert to the former dominant con- expenses incurred by conventional control mea- dition (Spies & Du Plessis 1987). sures have fostered the hope that biological con- Chemical and mechanical control of lantana was trol may provide a solution. reviewed by Cilliers & Neser (1991) who con- The biological control programme, including cluded that, although very effective, these meth- previously introduced natural enemies on L. ods are often labour-intensive and expensive. camara in South Africa, was reviewed by Cilliers & These control measures usually offer only tempo- Neser (1991), while other aspects of the weed were rary relief unless continual follow-up treatments reviewed by Swarbrick et al. (1995) and Munir are used. Controlled low- to moderate-intensity (1996). In this paper we mainly review biocontrol fires appear to reduce invasions by lantana, and initiatives undertaken during the 1990s, notably (i) can be an effective, preventative management the status of the natural enemies previously estab- Baars & Neser: Biological control of Lantana camara 23 Distribution of Lantana camara in South Africa. (Drawn by L. Henderson, Plant Protection Research Institute, Pretoria.) lished on lantana, (ii) the factors that have influ- The third species, the noctuid moth Neogalea enced the efficacy of these natural enemies, (iii) sunia (Guenée) (= N. esula (Druce), failed to estab- new biocontrol candidates under evaluation for lish despite introductions from Trinidad in 1962 release in South Africa and (iv) the problem of and 1968 and later from Australia in 1983 (Cilliers expanded host ranges of natural enemies under & Neser 1991). The remaining two species, Salbia laboratory conditions. haemorrhoidalis Guenée and Teleonemia scrupulosa Stål became established following the initial re- STATUS OF NATURAL ENEMIES leases (Table 2). ESTABLISHED IN SOUTH AFRICA Although widely established in South Africa, the Biological control of lantana in South Africa was flower-feeding pyralid, S. haemorrhoidalis, occurs initiated during 1961/62, with the introduction of mostly in low numbers and its impact on the weed five natural enemy species (Oosthuizen 1964). is unknown. By contrast, the sap-sucking lace bug, However, two of these, Hypena laceratalis Walker T. scrupulosa, has become widely established in (= H. strigata (F.)) and Ophiomyia lantanae high numbers throughout the range of L. camara in (Froggatt), were already established in South Af- South Africa. It is uncertain whether the later im- rica (Table 1) and were misidentified during a portation and release of additional genetic mate- pre-introductory survey conducted by Oost- rial from various countries (Table 2) has influenced huizen (1964). The leaf-feeding moth Hypena the efficacy of the tingid in South Africa (Cilliers & jussalis Walker was subsequently found to be syn- Neser 1991). Feeding damage is often extensive, onymous with H. laceratalis, while specimens of O. causing periodic defoliation of L. camara popula- lantanae were misidentified as O. rhodesiensis tions, with damaged plants suffering reductions in Spencer, which had been described from Zimba- seed set (Cilliers 1983, 1987a). Tingid populations bwe (Cilliers & Neser 1991). Hypena laceratalis is and their resultant damage peak in mid-summer widespread and damages seedlings and new but rapidly decline towards winter. Teleonemia growth in summer (Table1). Ophiomyia lantanae is scrupulosa is considered to be the most effective widespread and abundant, but has little impact on natural enemy of lantana in South Africa at present. seed viability (Cilliers & Neser 1991) (Table 1). Two leaf-mining hispine beetles, Octotoma 24 African Entomology Memoir No. 1 (1999): 21–33 Status of natural enemies present on Lantana camara in South Africa prior to the initiation of the biological control programme (‘generalists’ not included). Order/Family Natural enemy species Mode of attack Status Limitations Diptera Agromyzidae Ophiomyia lantanae (Froggatt) Fruit miner Widely established and abundant Low impact on seed viability; heavy parasitism Lepidoptera Noctuidae Hypena laceratalis1 Walker Leaf chewer Widely established with considerable Larvae are only active during late damage to seedlings and new growth. summer and autumn and are often Also attacks native Lippia and Priva parasitized species Pterophoridae Lantanophaga pusillidactyla1 Flower, fruit and Widely established, but present in low Low abundance and possible (Walker) seed chewer numbers. Also attacks native Lippia high levels of parasitism species Tortricidae Epinotia lantana1 (Busck) Flower-peduncle Widely established Unknown and shoot-tip borer Gracillariidae Cremastobombycia lantanella1 Leaf miner Widely established, but present in
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