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Crop Protection Compendium - Lantana camara L. Pierre Binggeli 2005 NAMES AND TAXONOMY Preferred scientific name Lantana camara L. Taxonomic position Domain: Eukaryota Kingdom: Viridiplantae Phylum: Spermatophyta Subphylum: Angiospermae Class: Dicotyledonae Order: Lamiales Family: Verbenaceae Other scientific names Lantana aculeata Lantana scabrida Ait. Lantana antidotalis Thonning (1827) Camara vulgaris Benth. Lantana camara var. aculeata BAYER code LANCA (Lantana camara) Common names English: lantana red-flowered sage white sage tickberry prickly lantana Spanish: venturosa mora de caballo cinco negritos cariaquillo camar corroncho comida de paloma santo negrito cinco cincos frutilla palo del diablo morita jarrila jaral soterre French: lantanier Cambodia: ach mann Fiji: kauboica Germany: wandelroeschen Guinea: boulé kogno kogno porto Haiti: bonbonier herbe au diable herbe bourrique herbe à plomb Hawaii: lakana India: bands nagaairi phullaki putus tantbi Indonesia: boenga pagar boenga pagar chente kembang satik kembang telek oblo puchengan puyengan saliara saliyere sliyara tahi agam tai hayam tai kotok telekan tembelek tembelekan teterapan waung wileran Kiribati: te kaibuaka Madagascar: fankatavinakoho fotatra mandadrieko radredreka rajejeka ramity Malaysia: bunga asam senyur bunga pagar bunga tahi anjing bunga tahi asu bunga tahi ayam busok Binggeli 2005 Crop Protection Compendium - Lantana camara L. 1 Assam: guphul Brazil: cambara de espinho bunga tahi ayam tahi ayam munai Mauritius: vieille fille Nicaragua: cuasquito Philippines: bahug-bahug sapinit Puerto Rico: cariaquillo Ryukyu Archipelago: shichi-henge Saint Helena: wild currant Samoa: Lantana latana South Africa: boesmandruiwe cherry-pie common lantana gewone lantana gomdagga sumba voelbrandewyn wild lantana wilderoosmaryn yellow sage Sri Lanka: ganda-pana garda-pana genda-pana katu-hinguru rata-guru ton-kinna Thailand: kamkung paka krawng pha-ka-krong Tonga: Talamoa talatala Vietnam: thom oi Zimbabwe: chiponiwe Notes on taxonomy and nomenclature L. camara is a highly variable species which has been widely cultivated for over 300 years. Hundreds of cultivars and hybrids exist (Howard, 1969) and most of them belong to the Lantana camara complex (Stirton, 1979). Cultivars can be distinguished morphologically (flower size, shape and colour; leaf size, hairiness and colour; stem thorniness; height and branch architecture), physiologically (growth rates, toxicity to livestock) and by their chromosome number and DNA content (Gujral and Vasudevan, 1983; Scott et al., 1997; Stirton, 1979). Two groups are often recognized: one with few or no spines commonly found in the neotropics and with spines in other parts of the world where the species is troublesome (Howard, 1970; Swarbrick, 1986). In the Pacific Islands the commonest variety is the prickly L. camara var. aculeata (Thaman, 1974). HOST RANGE List of hosts plants Major hosts Ananas comosus (pineapple), Camellia sinensis (tea), Cocos nucifera (coconut), Coffea (coffee), Durio zibethinus (durian), Elaeis guineensis (African oil palm), Gossypium (cotton), Hevea brasiliensis (rubber), Musa x paradisiaca (plantain), Oryza sativa (rice), pastures , Poaceae (grasses), Saccharum officinarum (sugarcane), Santalum album (Indian sandalwood), Shorea robusta (sal) HABITAT Habitat descriptors Principal weed in: managed forests; natural forests; wastelands Binggeli 2005 Crop Protection Compendium - Lantana camara L. 2 GEOGRAPHIC DISTRIBUTION Notes on distribution L. camara is native to Central and South America but its original distribution is unclear due to the introduction of a number of ornamental varieties. The species has also been poorly investigated in its native range, where it is not usually considered to be a serious pest, and the extent of its original native range is unclear. In the West Indies it is found in dry thickets (Adams, 1976). The weed is noted to be present in the Galapagos Islands of Ecuador (Cruz et al., 1986). Distribution List Europe Europe (as a whole) present Italy [Portugal] Azores Madeira present introduced (1692) introduced localized localized introduced introduced Spain localized introduced Asia Bangladesh present introduced invasive Brunei Darussalam present introduced invasive Cambodia China present present present, few occurrences widespread introduced introduced introduced (ca 1851) introduced introduced (1809) introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced invasive invasive Islam et al., 2001 Holm et al., 1979; Waterhouse, 1993 Waterhouse, 1993 Holm et al., 1979; Corlett, 1992 invasive Holm et al., 1979 invasive Xie Yan et al., 2001 invasive Burkill, 1935 invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive Rawat, 1997 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Gujral & Vasudevan, 1983 Sinha & Sharma, 1984 Sinha & Sharma, 1984 Gujral & Vasudevan, 1983 Nair & Henry, 1983 Gujral & Vasudevan, 1983 Hong Kong Taiwan India Andhra Pradesh Assam Bihar Delhi Himachal Pradesh Indian Punjab Jammu and Kashmir Karnataka Madhya Pradesh Maharashtra Orissa Rajasthan Tamil Nadu Uttar Pradesh present present widespread widespread widespread widespread widespread widespread widespread widespread present present widespread widespread widespread Binggeli 2005 Crop Protection Compendium - Lantana camara L. Burkill, 1935 Tutin et al., 1972 invasive not invasive Tutin et al., 1972 Press & Short, 1994 Sobrino et al., 2002 3 West Bengal Indonesia Java Kalimantan Sulawesi Israel present present widespread widespread widespread present, few occurrences introduced introduced introduced introduced introduced invasive invasive invasive invasive invasive introduced Sinha & Sharma, 1984 Waterhouse, 1993 Smiet, 1992 Holm et al., 1979 Whitten et al., 2002 Holm et al., 1979 [Japan] Ryukyu Archipelago localized introduced invasive Malaysia localized introduced invasive Myanmar present invasive Philippines widespread Saudi Arabia present introduced introduced (ca 1840) introduced Singapore present introduced invasive Sri Lanka widespread introduced (ca 1826) invasive Thailand present introduced invasive Timor-Leste Turkey widespread widespread introduced introduced invasive invasive Vietnam present introduced invasive Africa Angola present invasive Day et al., 2003 Cape Verde present invasive Chevalier, 1935 Comoros Congo Democratic Republic Côte d'Ivoire widespread introduced introduced (ca 1851) introduced invasive Roby & Dossar, 2000 present introduced invasive Day et al., 2003 present introduced invasive Ethiopia widespread introduced invasive Gabon Gambia Ghana Guinea Kenya Liberia present present widespread present widespread present introduced introduced introduced introduced introduced introduced invasive invasive invasive invasive invasive invasive Madagascar widespread introduced invasive Mauritius Rodriguez Island widespread widespread introduced introduced invasive invasive Holm et al., 1979 Binggeli & Desalegn Dessissa, 2002 Barreto et al., 1995 Robinson, 2001 Holm et al., 1979 Schnell, 1950 Holm et al., 1979 Holm et al., 1979 Holm et al., 1979; Binggeli, 2003 Macdonald et al., 1991 Strahm, 1989 Binggeli 2005 Crop Protection Compendium - Lantana camara L. invasive Walker, 1976 Holm et al., 1979; Waterhouse, 1993 Waterhouse, 1993 Burkill, 1935; Holm et al., 1979; Waterhouse, 1993 Day et al., 2003 Waterhouse, 1993; Baretto et al., 1995 Morton, 1994; Evans, 1999 Holm et al., 1979; Waterhouse, 1993 McWilliam, 2000 Holm et al., 1979 Holm et al., 1979; Waterhouse, 1993 4 Mozambique Namibia Nigeria Saint Helena Senegal Seychelles South Africa Sudan Swaziland Tanzania Uganda Zambia Zimbabwe Central America & Caribbean Antigua and Barbuda widespread present widespread widespread present present widespread present present widespread widespread present widespread introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced introduced present native Barbados widespread native Costa Rica Cuba Dominican Republic El Salvador Guatemala Haiti Honduras Jamaica Nicaragua Panama Puerto Rico Trinidad and Tobago United States Virgin Islands North America [USA] Alabama California Georgia (USA) widespread present present present present present present widespread widespread present localized widespread native native native native native native native native native native native native Francis et al., 1994 Gooding et al., 1965; Morton, 1994 Schemske, 1983 Sharma et al., 1988 Barreto et al., 1995 Day et al., 2003 Holm et al., 1979 Day et al., 2003 Holm et al., 1979 Adams, 1976 Holm et al., 1979 Holm et al., 1979 Holm et al., 1979 Holm et al., 1979 present native Morton, 1994 present present present introduced introduced introduced introduced (1859) introduced introduced introduced introduced introduced Doren et al., 2002 Morton, 1994 Doren et al., 2002 Hawaii widespread Louisiana Mississippi North Carolina Oklahoma South Carolina present present present present present Binggeli 2005 Crop Protection Compendium - Lantana camara L. invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive invasive Holm et al., 1979 Bromilow, 1995 Holm et al., 1979 Cronk, 1989 Holm et al., 1979 Gerlach, 1993 Erasmus et al., 1993 Barreto et al., 1995 Robertson et al., 2001 Holm et al., 1979 Holm et al., 1979 Holm et al., 1979 Holm et al., 1979 Davis et al., 1992 Doren et al., 2002 Doren et al., 2002 Morton, 1994 Doren et al., 2002 Doren et al., 2002 5 Texas Utah South America Argentina Bolivia [Brazil] Alagoas Amazonas Bahia Ceara Espirito Santo Goias Maranhao Matto Grosso do Sul Matto Grosso Minas Gerais Paraiba Parana Pará Pernambuco Piauí Rio Grande do Norte Rio Grande do Sul Rio de Janeiro Santa Catarina Sao Paulo Sergipe Colombia Ecuador Galapagos Islands Peru Venezuela Oceania American Samoa present present introduced introduced Morton, 1994 Doren et al., 2002 present present native native Morton, 1994 Holm et al., 1979 present present present present present present present present present present present present present present present present present present present present present present present widespread present present native native native native native native native native native native native native native native native native native native native native native native native introduced native native Lorenzi, 1983 Barreto et al., 1995 Barreto et al., 1995 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzo, 1983 Sharma et al., 1988 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Barreto et al., 1995 Lorenzi, 1983 Lorenzi, 1983 Lorenzi, 1983 Morton, 1994 Barreto et al., 1995 Cruz et al., 1986 Morton, 1994 Morton, 1994 widespread invasive Thaman, 1974 Australia widespread introduced introduced (1841) invasive Swarbrick, 1986 introduced invasive Swarbrick, 1986 introduced introduced invasive invasive not invasive not invasive Swarbrick, 1986 Swarbrick, 1986 Australian Northern localized Territory New South Wales widespread Queensland widespread South Australia localized introduced Victoria localized introduced Binggeli 2005 Crop Protection Compendium - Lantana camara L. invasive Swarbrick, 1986 Swarbrick, 1986 6 Western Australia Belau Cook Islands Federated states of Micronesia Caroline Islands Fiji localized widespread widespread introduced introduced introduced invasive invasive invasive Swarbrick, 1986 Meyer, 2000 Thaman, 1974 widespread introduced invasive Meyer, 2000 widespread widespread invasive invasive Thaman, 1974 Mune & Parham, 1967 French Polynesia widespread invasive Meyer, 2000 Guam Nauru localized widespread invasive invasive Thaman, 1974 Meyer, 2000 New Caledonia widespread invasive Heckel, 1911 New Zealand Niue Norfolk Island Northern Mariana Islands Papua New Guinea Pitcairn Islands Samoa Solomon Islands Tonga Vanuatu widespread widespread widespread introduced introduced introduced (1843) introduced introduced introduced (ca 1883) introduced introduced introduced invasive invasive invasive Holm et al., 1979 Meyer, 2000 Swarbrick, 1986 widespread introduced invasive Meyer, 2000 widespread widespread widespread present widespread widespread introduced introduced introduced introduced introduced introduced invasive invasive invasive invasive invasive invasive Holm et al., 1979 Florence et al., 1995 Thaman, 1974 Swarbrick, 1986 Thaman, 1974 Mullen et al., 1993 HISTORY OF INTRODUCTION AND SPREAD This ornamental shrub has been introduced throughout the tropics and subtropics, often used as a hedge plant, and is commonly grown in the temperate zone. It can grow between the latitudes 45°N and 45°S and an altitude of up to 1400 m. Although first cultivated in Europe during the late 17th century and reached Calcutta in 1809 (Burkill, 1935). It was mostly introduced throughout the tropics during the later part of the 19th century and a number of cultivars and forms were subsequently disseminated (Howard, 1970). In the tropics it is a weed of a number of crops and in many countries it has been declared a noxious plant. In many tropical regions the thorny forms have invaded huge areas of natural pasture land. In Singapore L. camara became for some time quite abundant but by around 1900 it became less noticeable (Burkill, 1935) and a similar phenomenon has been reported for East Timor (McWilliam, 2000). BIOLOGY AND ECOLOGY Genetics The known chromosome numbers are 2n = 22, 33, 44, 55, but most invasive varieties of L. camara appear to be tetraploids (Day et al., 2003). Besides variation in chromosome number there is much variation in DNA content, growth rates and toxicity to livestock Binggeli 2005 Crop Protection Compendium - Lantana camara L. 7 (Stirton, 1979; Gujral and Vasudevan, 1983; Scott et al., 1997). In the Tamil Nadu region of India there are differences in toxicity of L. camara, with the red flowered variety being more toxic than the pink flowered form (Thirunavukkarasu et al., 2001). Physiology and Phenology Flowering and fruiting take place throughout the year with a peak during the first 2 months of the rainy season. In the highlands of western Kenya an investigation of leaf decomposition found that after 7 days it had decreased to just under a third of the original mass and by the 77th day the leaves had totally decomposed. The percentage of the initial amount of phosphorus and nitrogen remaining in the leaf material after a week was 42 and 54%, respectively. After 21 days 90% of the phosphorus had been released (Kwabiah et al., 2001). Reproductive Biology The flowers of L. camara, when yellow, produce nectar and are pollinated by butterflies and thrips. The species is an obligate outcrosser and it is unclear whether apomixis occurs. Fruits mature rapidly and change colour from dark green to black. A number of bird species, and also sheep and goats disperse the seeds, sometimes over long distances, but natural dispersal between oceanic islands has never been demonstrated. Heavy fruit crops are produced yearly, but the thornless forms produce few, if any, seeds. Seeds germinate when sufficient moisture is available, usually at the start of a rainy season. In Australia, Broughton (1999) found that 57-80% of green and ripe fruits tested had one or two viable seeds whereas 12 and 34% had none, and between 64 and 90% of dried (older) fruits had two nonviable embryos suggesting that fruit development stage affects germination. She found no difference in viability within sites or between cultivars investigated. In addition to spreading by seed, L. camara is able to produce adventitious shoots, especially shallow lateral roots, following mechanical damage. Hence, it is also able to spread and establish dense thickets by vegetative means. The capacity of the species to spread vegetatively and to inhibit both the growth of other vegetation and seed germination, in conjunction with heavy and regular fruiting, is the main reason why L. camara forms long-lasting permanent thickets. In areas where natural fires occur they stimulate thicker regrowth. For further information, see Mathur and Mohan Ram (1978), Schemske (1983), Sinha and Sharma (1984) and Thaman (1974). Environmental Requirements L. camara can grow between the latitudes 45°N and 45°S and an altitude of up to 1400 m. The rapid spread of L. camara throughout the tropics is associated with human-induced disturbances. It forms extensive, dense and impenetrable thickets in forestry plantations, orchards, pasture land, waste land and in natural areas. L. camara thrives in open and disturbed areas as well as in open natural vegetation. Being somewhat shade-tolerant it can become the dominant understorey shrub in open forests, but is absent from closed forests. L. camara grows under a wide range of climatic conditions. In Australia it tolerates a mean annual rainfall from 4000 to less than 1000 mm, and as low as 200 mm per annum elsewhere (Gujral and Vasudevan, 1983). It is found between sea level and nearly 1000 m on Hawaii, and higher in East Africa, the upper altitudinal limit being determined by frost, which the plant is susceptible to. In Hong Kong, temperature in the range 3-5°C injured L. camara (Corlett, 1992). It tolerates salt spray. Its distribution is affected by soil type. It has a low tolerance for boggy and saline soils but grows well on poor soils. Binggeli 2005 Crop Protection Compendium - Lantana camara L. 8 L. camara has a marked ability to compensate for herbivory as plants survived experimental defoliation for 2 years (Broughton, 2000). Associations L. camara often occurs in pure stands but can be mingled with a variety of species but emergent shrubs and trees in particular. Climatic amplitude (estimates) - Mean annual rainfall: 200 - 4000 mm - Rainfall regime: summer; bimodal - Mean annual temperature: 13 - 0ºC Soil descriptors - Soil texture: medium; heavy - Soil drainage: free - Soil reaction: acid; neutral - Special soil tolerances: infertile MEANS OF MOVEMENT AND DISPERSAL Natural Dispersal (Non-Biotic) Occasionally abiotic seed dispersal may occur. Flash floods in South Africa, caused by cyclone Demoina in 1983, transported seeds and deposited them on the flood plain of the Ndumu game reserve (Bromilow, 1995). Vector Transmission (Biotic) The seeds are dispersed by native or invasive species of birds. In Hong Kong, L. camara is dispersed by 15 species of native birds (Corlett, 1998), whereas in Hawaii, it is mainly dispersed by exotics such as the Indian myna (Atkinson and Atkinson, 2000). Accidental Introduction Accidental introduction via contaminated soil is possible but has not been documented. Intentional Introduction As L. camara is such a key ornamental plant, new varieties, some of which have invasive potential, can readily be bought and introduced throughout the tropics. Transport pathways for long distance movement - Soil, Gravel, Water, Etc. NATURAL ENEMIES The alkaloid-rich leaves of L. camara are virtually immune to grazing by livestock, although several hundred phytophagous insects have been recorded on it. In the New World flowers, flower stalks, leaves, shoots and roots are attacked by many insect species and pathogens although their impact on shrub vigour and seed set is poorly understood. In Mexico a stem Binggeli 2005 Crop Protection Compendium - Lantana camara L. 9 sap-sucking membracid bug, Aconophora compressa, causes considerable dieback of stems (Swarbrick et al. 1995). Elsewhere there are no reports of important pests and diseases besides the introduced biocontrol agents. The following fungi have been found attacking the leaves of L. camara: Dendryphiella aspera, Micropustulomyces mucilaginosus, Mycovellosiella lantanae var. lantanae, Septoria sp., Ceratobasidium lantanae, Prospodium tuberculatum and Puccinia lantanae. For further information on fungal natural enemies of L. camara, see Barreto et al. (1995), Den Breeyen et al. (2000), Thomas and Ellison (2000), Trujillo and Norman (1995). Natural enemies listed in the database The list of natural enemies has been reviewed by a biocontrol specialist and is limited to those that have a major impact on pest numbers or have been used in biological control attempts; generalists and crop pests are excluded. For further information and reference sources, see About the data. Additional natural enemy records derived from data mining are presented as a separate list. Natural enemies reviewed by biocontrol specialist Natural enemy Pest stage attacked Biological control in: Herbivores: Aceria lantanae Inflorescence, Leaves Aconophora compressa Leaves Aerenicopsis championi Stems Alagoasa prob. quadrilineata Leaves Anoncia diveni Leaves Autoplusia illustrata Leaves Calycomyza lantanae Leaves Charidotis pygmaea Leaves Coelocephalapion Inflorescence, Leaves Cremastobombycia lantanella Leaves Diastema tigris Leaves Ectaga garcia Leaves Crocidosema lantana Eutreta xanthochaeta (gall fly, lantana) Fruits/pods Geraeus nr. curvispinis Inflorescence Hepialus Stems Hypena laceratalis Leaves Langsdorfia franckii Roots Lantanophaga pusillodactyla Inflorescence Leptobyrsa decora Leaves Longitarsus Leaves Neogalea sunia (lantana stick, caterpillar Leaves (USA)) Octotoma championi Leaves Octotoma pliculata Leaves Octotoma scabripennis (leaf beetle, lantana) Leaves Omophoita albicollis Inflorescence, Leaves Ophiomyia camarae Leaves Ophiomyia lantanae (fly, lantana seed) Fruits/pods, Inflorescence Plagiohammus spinipennis Stems Pseudopyrausta acutangularis Leaves Binggeli 2005 Crop Protection Compendium - Lantana camara L. Asia Micronesia Hawaii Micronesia New Caledonia; South Africa Asia; Micronesia; South Africa 10 Salbia haemorrhoidalis Strymon bazochii Teleonemia elata Teleonemia harleyi Teleonemia prolixa Teleonemia scrupulosa (lantana, bug) Teleonemia validicornis Thecla Strymon bazochii Uroplata girardi (lantana, hispid) Leaves Fruits/pods, Inflorescence Leaves Leaves Leaves Leaves Leaves Leaves Leaves Leaves Uroplata lantanae Leaves South Africa Micronesia; South Africa Micronesia; New Caledonia; South Africa Additional natural enemies (source - data mining) Natural enemy Pest stage attacked Biological control in: Herbivores: Hypena strigatus (lantana defoliator, caterpillar) Platyptilia pussilidactyla South Africa IMPACT Economic impact In Central America L. camara is common in pastures, waste areas and roadsides; it is also a weed in a number of crops (Schemske, 1983), although infestations are unlikely to be composed of native biotypes, but rather re-introduced cultivars that have become invasive (Stirton, 1977). In many countries L. camara encroaches on agricultural land, reduces the carrying capacity of pastures and is a weed in many agricultural crops. In Australia, L. camara has infested about 4 million ha of pasture (Parsons and Cuthbertson, 1992). In the early 1980s this resulted in economic losses of A$7.7 m (Swarbrick et al., 1995). Holm et al. (1977) reported that in some areas of India the invasion of cultivated lands by this weed led to the shifting of several villages. In Fiji it is a major weed of coconut plantations, pastures, neglected arable land and waste places (Mune and Parham, 1967). In forestry it tends to over-run young plantations, prevent access to older ones and increase fire hazards. In Indian sandalwood forests the shrub competes with sandalwood trees and also favours the spread of the sandal spike disease. In contrast to the widely held view that L. camara is detrimental, Timorese farmers have considered the plant as highly beneficial as it enhanced soil fertility and soil conditioning. This resulted in a reduction in fallow periods under L. camara from 15 to 5 or 6 years. Another benefit was the supply of firewood (McWilliam, 2000). The idea that L. camara enhances soil fertility has yet to be demonstrated and Binggeli (2001) has postulated that the Pitcairners' selection of sites with thriving L. camara stands for home gardens reflects the species predilection for fertile sites rather than its ability to increase fertility. Binggeli 2005 Crop Protection Compendium - Lantana camara L. 11 Environmental impact In natural and semi-natural vegetation L. camara is a major conservation problem. It may smother vegetation and increase fire intensity (due to an increase in dry biomass), thus displacing native scrub communities (e.g. Heckel, 1911). Its extensive seed production favours rat populations. There are many unsubstantiated statements suggesting that L. camara slows erosion (e.g. Ashmole and Ashmole 2000), but it is likely that this may be the case when the plant becomes established on bare ground but not when it displaces native vegetation. It can grow through the pestiferous grass Imperata cylindrica and suppress it in South-East Asia and thus has some potential in forest restoration (Burkill, 1935). Social impact Stands of lantana, and of the prickly variety in particular, hinder human's access to invaded habitats. In Tanzania and Uganda, L. camara can be considered a serious health hazard, as its thickets provide breeding grounds for tsetse flies, vectors of trypanosomiasis (Leak, 1999). L. camara thickets are potential breeding places for rats, wild pigs, insect pests and plant diseases. When ingested by cattle and sheep it may cause photosensitive reactions, diarrhoea, jaundice, hepatitis and poisoning. Children have been known to die after eating unripe berries and stems have been used as for toothbrushes (Burkill, 1935; Morton, 1994; Swarbrick et al., 1995). Impact on biodiversity L. camara can readily hybridize with other Lantana species, for example, in Florida it hybridizes with the endangered endemic L. depressa (Langeland and Burks, 2000). The impact on native vegetation is mainly viewed as negative, i.e. reducing species diversity, threatening endemics (Cruz et al., 1986) and leading species to extinction. In Australia, L. camara causes allelopathic suppression of two indigenous tree species (Gentle and Duggin, 1997). It is also generally considered to hinder the regeneration of native tree species (e.g. Islam et al., 2001) but there are some occasional references to regeneration of some tree species under its canopy (e.g. Burkill, 1935). The spread of L. camara on the Galapagos Islands is seen as a threat to bird breeding populations (Cruz et al., 1986). The impact of L. camara on biodiversity is mostly negative but a few instances of a positive impact have been reported. It is often said that it provides habitat for some birds and thus provides refuge for wildlife (Mullen et al., 1993). More specifically, in Kenya lantana thickets have been reported to harbour a threatened bird species (Hinde's Babbler, Turdoides hinduei). It provides shelter to the bird that is not now readily available in a humandominated countryside (Njoroge and Bennun, 2000). The plant plays a minor role in the feeding ecology of some species of conservation interest such as the lion-tailed macaque (Macaca silenus), which feeds extensively on the fruits in southern India (Umapathy and Kumar, 2000). As it is such a variable species, including variability in stature, specific varieties or forms can be expected to have different impacts on native biodiversity, as well as cropping systems and other human activities; however, no information is available regarding these potential differences. For more information see Holm et al. (1977), Morton (1994), Schemske (1983), Sharma et al. (1988), Sinha and Sharma (1984) and Thaman (1974). Binggeli 2005 Crop Protection Compendium - Lantana camara L. 12 Summary of impact Negative impact on: biodiversity; environment; livestock production; forestry production; human health; native fauna; native flora; transport and travel; tourism Negative impact on: crop production PHYTOSANITARY SIGNIFICANCE The horticultural trade, in particular the availability of numerous varieties on the internet, is the main risk for the introduction of weedy varieties of L. camara. SUMMARY OF INVASIVENESS L. camara is a highly variable ornamental shrub, native of the neotropics. It has been introduced to most of the tropics and subtropics and is now extremely weedy in many countries. It is generally deleterious to biodiversity and human activities and has been the target of control programmes for a century; however, successful control has only been achieved in a few instances using biocontrol agents. Risk and Impact Factors • • • • • • • • • • • • • • • • • • • • • invasive in its native range: no proved to be invasive outside its native range: yes highly adaptable to different environments: yes high reproductive potential: yes highly mobile locally: yes its propagules remain viable for more than one year: unknown tolerates cultivation, browsing pressure, mutilation, fire etc.: yes competitive in crops or pasture: yes affects ecosystem: yes adversely affects natural communities: yes adversely affects community structure: yes adversely affect human health: no has sociological impacts on recreational patterns, aesthetics, property values: unknown harmful to animals: yes produces spines, thorns or burrs: yes host or vector of pests or diseases: yes likely to be accidentally transported internationally: yes likely to be deliberately transported internationally: yes difficult to identify or detect as a commodity contaminant: yes difficult to identify or detect in the field: no difficult or costly to control: yes MORPHOLOGY Plant type: ; succulent; woody; seed propagated; perennial. L. camara is a perennial shrub 2-5 m tall. It is erect in open areas whereas in scrubland it is scrambling. The stem is 4-angled, often bearing recurved prickles. The serrated leaves Binggeli 2005 Crop Protection Compendium - Lantana camara L. 13 are ovate to ovate-lanceolate (up to 10 cm long and 7.5 cm wide) and usually light coloured. The plant has a strong, unpleasant odour. Flowers (9 mm long) are usually yellow and pink, later turning orange then red and sometimes blue or purple. The black fleshy drupes, borne in clusters, are 3-6 mm in diameter and contain 1-2 seeds (1.5 mm long). SIMILARITIES TO OTHER SPECIES Although large stands of weedy varieties of L. camara are easily recognized, it is in fact a variable polyploid complex of interbreeding taxa resulting from hybridization with species in the other complexes, such as L. urticifolia (Day et al., 2003). In Florida, USA, it may be confused with the endangered endemic native, Lantana depressa, with which it has extensively hybridized (Langeland and Burks, 2000). DETECTION AND INSPECTION L. camara is conspicuous due to its attractive and multicoloured floral displays, and is wellknown throughout the tropics. CONTROL Cultural Control Being poisonous to stock means that the species can not be controlled using large herbivores. In fact, intense grazing by goats and donkeys will favour L. camara infestations by suppressing competition from palatable species (Ashmole and Ashmole, 2000). Mechanical Control Mechanical control can be effective, particularly where land is cleared, but requires continual follow-up treatment to remove roots and seedlings of L. camara. Slashing and burning stimulate suckering. Both chemical and mechanical control methods are expensive and labour intensive and are only effective in the short term. Cleared areas are rapidly colonized via seeds originating from distant parents or from sprouting roots. Chemical Control The Australian experience in controlling L. camara, reviewed by Swarbrick et al. (1995), indicates that some herbicides are more effective on particular lantana forms. The most effective herbicides belong to the phenoxy acid (2,4-D, 2,4,5-T, dichloroprop and MCPA), benzoic acid (dicamba) and pyridine groups. Glyphosat, sulfonylureas (metsulfuron methyl) and imidazolinones (imazapyr) also show good activity. Photosynthetic herbicides (triazine and urea) are not effective. A number of factors affect the effectiveness of the chemical treatment and they include: plant size, time of application, mode of application, and the use of surfactant. Use of herbicide in uncut stands may not be effective in preventing eventual regrowth. Combination of mechanical and chemical control may be the best. The seasonal response of L. camara to applications of fluroxypyr, metsulfuron-methyl, glyphosate and dichlorprop has recently been reported by Hannan-Jones (1998). Recent work carried out in the South African Kruger National Park by Erasmus et al. (1993) showed that chemical control was cheaper and caused less disturbance resulting in higher Binggeli 2005 Crop Protection Compendium - Lantana camara L. 14 biodiversity than mechanical control. Chemical control consisted in an application of imazapyr on freshly cut stems and a follow-up operation by spot-spray application of glyphosate. The initial control required 25 man-days per ha and that of the follow-up control 6.8 man-days per ha. Control costs will vary from site to site and will depend on L. camara stem density and cover. Latest South African recommendations are provided by Vermeulen et al. (1996). In India, eradication of L. camara from sub-watersheds in the Markanda catchment, Himal Pradesh, was effective and economical using glyphosate sprayed on to regenerated growth, cut 4 months previously (Rana and Singh, 1999). L. camara is resistant to triclopyr, a widely used herbicide for woody weed control (Goodall and Naude, 1998). Biological Control Since the turn of the century, biological control of L. camara has been attempted in many parts of the tropics with varying degrees of success, as different cultivars display differences in susceptibility to insect herbivores (Julien and Griffiths, 1998). Worldwide well over 200 releases have been made (39 different natural enemies have been released in 29 countries), however, in the majority of cases the control agent either failed to become established or became established without achieving control. Most of the releases have been carried out in the Pacific, South Africa and Australia (for historical details see Taylor 1989; Cilliers and Neser, 1991; Davis et al., 1992; Denton et al., 1991; Swarbrick et al., 1995). The most widely established species include Ophiomyia lantanae, Uroplata girardi and Octoma scabripennis. Day et al. (2003) have produced a detailed review of 48 of these control agents. In Hawaii, Neogalea sunia and Epinotia lantanae contribute usefully to the control of lantana across the islands. In addition, a combination of Hypena strigata, Octotoma scabripennis, Salbia haemorrhoidalis, Teleonemia scrupulosa and Uroplata girardi provide partial to substantial control in drier areas (<1270 mm rainfall), and in wetter areas Plagiohammus spinipennis provides partial control (Julien and Griffiths, 1998). The release in 1993 of Uroplata girardi on an island of the Russell Island group (Solomon Islands) resulted in the successful control of the 'Hawaiian Pink' form (Swarbrick et al., 1995). U. girardi has proved to be one of the more successful agents and is credited with providing some check on the spread of lantana in Australia, South Africa and some islands in the Pacific Ocean. In Micronesia seven out of 13 introduced insect species became established and have resulted in acceptable levels of control for current agricultural practices (Denton et al., 1991). As elsewhere the effectiveness of various insect species varied between islands and between lantana varieties. Greater success appears to have been achieved in drier areas. In Uganda, the introduction of Teleonemia scrupulosa, which had been widely released after its successful introduction into Hawaii in 1902, was very successful in the area around Serere Research Station in Teso District but it also attacked one of the cultivars of Sesamum indicum grown on the Research Station (Davies and Greathead, 1967). Fortunately, it was unable to breed on that crop and attacks subsided after the lantana had been controlled. Subsequently other agents for lantana control were tested on Sesamum and it was found that other Tingidae and the chrysomelid leaf miners would also feed on this crop (Greathead, 1973). The polyphagous pest Phenacoccus parvus severely damages stands of L. camara in Australia and is not, as commonly reported, a pest of potato and aubergine (Solanum melongena), although it has the potential to attack a variety of plant Binggeli 2005 Crop Protection Compendium - Lantana camara L. 15 species inclusive of some crops (Marohasy, 1994). A century of efforts has resulted in the various biocontrol agents providing partial to substantial success in controlling L. camara (Broughton, 2000). Despite this limited success, classical biological control is still considered to be the only viable, long-term control option, since it offers a safe, economic and environmentally benign method of suppressing the weed. New control agents are being actively sought in Mexico and South America and include both insects and fungal pathogens (Barreto et al., 1995). In South Africa it is hoped that a better understanding of the functional relationships and interactions between L. camara cultivars and insect herbivores may enhance the prospects of successful biological control (Cilliers and Neser, 1991). A potential control agent of lantana (spreading in Hawaii) has recently been identified by Trujillo and Norman (1995). It is a leaf spot fungus, Septoria sp., from Ecuador and is presumed to be potentially effective in controlling L. camara. In South Africa, the fungus Mycovellosiella lantanae var. lantanae, collected from Florida, USA, is soon to be released (Den Breeyen et al., 2000); and in Australia, permission has just been granted by the quarantine authorities to import the rust fungus Prospodium tuberculatum (ex Brazil). Other pathogens with apparently excellent potential to control a wide range of cultivars have been identified (Barreto et al., 1995; Thomas and Ellison, 2000). Broughton (2000) reviewed biological control programmes of L. camara worldwide and concluded that leaf-, flower- and fruit-feeding species were the most successful feeding groups, and the leaf-mining chrysomelid Uroplata girardi was the most successful control agent. She identified the main factor preventing the establishment of control agents as the number of individuals released and noted that cultivar preferences, parasitism and predation, and climate reduced control. Broughton (2000) concluded that flower- and fruitfeeding species were unlikely to be effective because the seeds of L. camara are only viable for a short period of time and have a low germination, and that defoliating species were likely to be ineffective because of the ability of L. camara to withstand defoliation. USES Since the 19th century L. camara has been one of the main tropical and subtropical garden ornamentals. Under temperate climes it has been, and still is, widely used as a glasshouse ornamental and a pot plant. Apart from its ornamental value, L. camara has few redeeming features. In some mountainous areas (e.g. in Tanzania and India) the presence of L. camara was once considered a good ground cover preventing erosion. In parts of East Africa, in locations where it is not weedy, it has effectively been used as a live fence (Howes, 1946). However, in parts of Ethiopia where the idea of establishing a live lantana fence to protect crops from domestic animals was taken up by local villagers in the 1990s, this quickly led to the loss of rough grazing land through the rapid spread of lantana (Binggeli and Desalegn Desissa, 2002). A number of minor uses of L. camara include using the seeds as a source of food for lambs, using lantana straw mixed with dung for biogas production, and using the twigs as fuel. There is some evidence, although conflicting in nature, that extracts from lantana may have value as biocides (Ahmed and Agnihotri, 1977). In addition, essential oils from the flowers and leaves may have some value to the perfume industry and as beneficial drugs (Ahmad et al., 1962). In parts of its native range, L. camara is used as a source of medicinal cures, for example, in Ecuador the leaves are ingested to treat stomach disorders (Ellison and Evans, 1996). It is viewed in many regions as an important honey plant (Fichtl and Admasu Adi, 1994). Leaf extracts have strong insecticidal and antimicrobial activity, for example, storing potatoes with lantana leaves almost eliminates damage by the potato Binggeli 2005 Crop Protection Compendium - Lantana camara L. 16 tuber moth Phthorimaea operculella (Lal, 1987). PESTS Pests listed in the database Major host of: Acherontia atropos (death's Head Hawkmoth), Omiodes indicata (soyabean webworm), Phenacoccus madeirensis (cassava mealybug) Minor host of: Acherontia styx (small death's head hawkmoth), Aphis gossypii (cotton aphid), Ectropis bhurmitra (tea twig caterpillar), Maconellicoccus hirsutus (pink hibiscus mealybug), Phytonemus pallidus (strawberry mite), Pratylenchus penetrans (nematode, northern root lesion), Pseudococcus jackbeardsleyi (Jack Beardsley mealybug) Wild host of: Eutypa lata (Eutypa dieback), Icerya purchasi (cottony cushion scale), Orthezia insignis (greenhouse orthezia), Piezodorus hybneri (legume stink bug), Rhizobium radiobacter (crown gall), Rhizobium rhizogenes (gall) Host of (source - data mining): Atta insularis , Phenacoccus parvus , Trialeurodes vaporariorum (whitefly, greenhouse) REFERENCES Adams CD, 1976. Flowering plants of Jamaica. Mona: University of the West Indies. Ahmad MN, Bhatty MK, Karimullah, 1962. 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