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Review and Evaluation of Lantana Biocontrol Programs

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Review and Evaluation of Lantana Biocontrol Programs Biological Control 17, 272–286 (2000) doi:10.1006/bcon.1999.0793, available online at http://www.idealibrary.com on COMMENTARY Review and Evaluation of Lantana Biocontrol Programs Sonya Broughton1 Department of Entomology and Zoology, The University of Queensland, St Lucia, Queensland 4072, Australia Received February 16, 1999; accepted October 8, 1999 feeding insect species were identified as successful This paper provides a review of lantana (Lantana agents, but different species have been successful in camara L.) biological control programs worldwide. different countries (Crawley, 1986, 1989a,b). Tables on the origins of the agents introduced for the Inability to predict success is attributed to the low biocontrol of lantana, are presented, including refer- genetic uniformity of lantana and its ability to colonize ences to the biology and/or host-tests for each species. diverse habitats (Crawley, 1989a,b; Willson, 1993; Swar- Establishment and control rates of the introduced agents and cases leading to partial control of lantana brick et al., 1995). Because of this, Willson (1993) are discussed. From the review, feeding groups and suggested that agent selection procedures proposed by species contributing to control were identified. Leaf-, Harris (1973a,b), Goeden (1983), and Wapshere (1985) flower-, and fruit-feeding species were the most success- are irrelevant. Instead, species were prioritized accord- ful feeding groups, and the leaf-mining chrysomelid, ing to their success in other countries or because they Uroplata girardi Pic, was the most successful control were host-specific and added to the folivore complex agent. The main factor preventing establishment was already introduced (Harley and Kassulke, 1971; Water- the number of individuals released, while cultivar house and Norris, 1987; Willson, 1993). However, this preferences, parasitism and predation, and climate selection method has failed to achieve successful con- reduced control. The implication of these results for trol, and many countries are introducing new species to lantana biocontrol programs is discussed, and future maximize lantana suppression (Willson, 1993; Neser, research requirements are identified. ௠ 2000 Academic Press Key Words: biological control; weeds; lantana; Lan- 1994; Palmer and Pullen, 1995; Swarbrick et al., 1995). tana camara; Uroplata; Octotoma; Teleonemia; Plagio- Because each new species costs 3 to 5 scientist-years in hammis. terms of research associated with host-specificity test- ing, development of mass-rearing techniques, and re- leases (Waterhouse and Norris, 1987; Harris, 1991), INTRODUCTION guidelines that improve selection of effective natural enemies are required. For almost a century, attempts have been made to The objective of this paper was to develop selection control Lantana camara (L.) using insects (Perkins and guidelines based on an analytic review of all lantana Swezey, 1924; Holloway, 1964; Swarbrick et al., 1995). biocontrol programs. Waterhouse and Norris (1987) To date, 38 species have been released in 29 countries provide the most complete review of lantana programs, (Willson, 1993; Swarbrick et al., 1995; Julien and but only for Pacific Rim countries. Neser and Cilliers Griffiths, 1998). Despite the amount of research effort (1989) supply some guidelines, but base their conclu- and time expended on lantana biological control, re- sions on the 1982 review of Julien (1982), or papers sults have been variable. Success in several countries published before 1987, and did not include statistical led Crawley (1986, 1989a,b) to rate lantana as the analysis of programs. Using a variation of the success ‘‘most successful target of weed biocontrol,’’ but he also rating of Crawley (1989a,b), I analyzed current data on listed lantana as the most ‘‘frequent unsuccessful tar- the success of individual lantana agents in controlling get’’ because of failures in many locations. Three leaf- lantana in every location of release. The following factors were included in my analyses to identify trends in agent selection or correlations with establishment or 1 Present address: Entomology Department, Agriculture Western Australia, 3 Baron-Hay Court, South Perth, Western Australia 6151, control of lantana: Lantana taxon from which the Australia. biocontrol agent had been collected, insect taxa, part of 1049-9644/00 $35.00 272 Copyright ௠ 2000 by Academic Press All rights of reproduction in any form reserved. REVIEW AND EVALUATION OF LANTANA BIOCONTROL PROGRAMS 273 the plant attacked, cultivar preference, number of 1; Spies, 1984; Swarbrick, 1985; Waterhouse and Nor- individuals released, and natural enemies. ris, 1987; Palmer and Pullen, 1995). Lantana taxon. Lantana biocontrol agents have Weed scientists often refer to cultivars as L. camara, been collected from many different species of Lantana implying that they all belong to one monotypic species. under the assumption that coevolutionary history or However, reports suggest that cultivars were derived ‘‘old’’ vs ‘‘new’’ insect–plant associations (Hokkanen and from different lantana species or subspecific taxa (e.g., Pimental, 1984) do not affect the likelihood of establish- Lantana camara aculeata and L. camara var. mista) ment or control. However, this premise was never and many are of indeterminate origin (Table 1). Culti- tested for lantana. vars have been classified according to flower color, Insect taxa. Harley (1971), Winder and Harley presence or absence of thorns, and growth habit (Smith (1982), and Neser and Cilliers (1989) suggested that and Smith, 1982). Cultivars differ in their susceptibil- Lepidoptera were less likely to control lantana because ity to insect attack, chemical constituents, and toxicity they usually are heavily parasitized. to livestock (Smith and Smith, 1982; Neser and Cilliers, 1989; Taylor, 1989; Cilliers and Neser, 1991). In most Part of the plant attacked. Crawley (1989a,b) identi- countries, a pink and yellow flower cultivar is common fied insects that attacked the leaves as the most (Table 1). Scott et al. (1997) recently questioned the use successful feeding group. Perkins and Swezey (1924) of flower color to classify lantana. Their comparison of considered flower- and fruit-feeding species to be the the most common cultivars in Queensland, Australia, most successful feeding group. revealed no correlation between flower color and culti- Cultivar preference. Preferences of agents for par- var based on DNA analysis. ticular cultivars were thought to limit control (Harley, 1973; Neser and Cilliers, 1989; Swarbrick et al., 1995). Number of individuals released. Beirne (1985) sug- CHARACTERISTICS OF INSECTS SELECTED gested that the chances of colonization were less than AS BIOCONTROL AGENTS 10% if 5000 or fewer individuals are released. The insect species surveyed worldwide were com- Natural enemies. Goeden and Louda (1976), Cullen pared with those released for lantana biocontrol on the and Snowball (1979), and Crawley (1989a,b) identified basis of insect order, part of plant attacked, and host natural enemies (predators, parasitoids, disease) as specificity. Survey data were obtained from Palmer and some of the most important factors that limit establish- Pullen (1995) and Winder and Harley (1983), and data ment and control of biological control agents. on released insects were from host-testing reports or These factors are discussed relative to the origin of from Swarbrick et al. (1995). lantana, the insects surveyed and those introduced for biocontrol, the success of all introduced agents in terms Worldwide Surveys of establishment and control of lantana, the factors that limit establishment and/or control, and the re- Potential biological control agents were surveyed in sponses of lantana to biocontrol agents. This informa- Brazil, Peru, Colombia, and Central and North America tion was used to derive guidelines for future programs. (Perkins and Swezey, 1924; Krauss, 1953, 1962; J. Mann, unpublished data, 1954; Harley, 1974; Diatloff, 1977; Winder, 1980; Winder and Harley, 1983; Palmer ORIGIN OF LANTANA and Pullen, 1995). Cumulatively, these surveys re- sulted in the collection of 550 phytophagous insect The specific geographic origin of L. camara is un- species from Lantana spp. in Central and North America known. Some authors suggested that L. camara is and 345 species from Brazil (Winder and Harley, 1983; native to South America or Mexico (Howard, 1969; Palmer and Pullen, 1995). Smith and Smith, 1982; Spies and du Plessis, 1987), while others suggested the West Indies as the site of Taxa and Host Range origin (Moldenke, 1973; Stirton, 1977; Palmer and Pullen, 1995). In the mid-16th and 17th centuries, Species were divided with respect to taxonomic lantana species, including L. camara, were imported groups, plant part attacked, and host specificity (Figs. 1 into Europe as horticultural plants from the Americas and 2). Polyphagous species are defined by Harley and (Stirton, 1977; Swarbrick et al., 1995). Hundreds of Forno (1992) as ‘‘insects that feed on many different cultivars of mixed parentage were created in Europe species of plants.’’ Oligophagous species feed on a single from this stock (Howard, 1969). These cultivars were order, family, or genus, while monophagous insects are transported back to the Americas and to Australia, restricted to a single plant species (Harley and Forno, India, and Africa in the mid-19th century (Table 1; 1992). Howard, 1969; Stirton, 1977; Swarbrick, 1985). Many Coleoptera, particularly Chrysomelidae, was the cultivars escaped
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