lantana camara l. - mikepalmerbunga tahi anjing bunga tahi asu bunga tahi ayam busok binggeli 2005...

Crop Protection Compendium - Lantana camara L.

Pierre Binggeli 2005

NAMES AND TAXONOMY

Preferred scientific name

Lantana camara L.

Taxonomic positionDomain: EukaryotaKingdom: ViridiplantaePhylum: SpermatophytaSubphylum: AngiospermaeClass: DicotyledonaeOrder: LamialesFamily: Verbenaceae

Other scientific namesLantana aculeataLantana scabrida Ait.Lantana antidotalis Thonning (1827)Camara vulgaris Benth.Lantana camara var. aculeata

BAYER codeLANCA (Lantana camara)

Common names

English: lantanared-flowered sagewhite sagetickberryprickly lantanaSpanish: venturosamora de caballocinco negritoscariaquillocamarcorronchocomida de palomasanto negritocinco cincosfrutillapalo del diablomoritajarrilajaralsoterreFrench: lantanier

Assam: guphulBrazil: cambara de espinho

Cambodia: ach mannFiji: kauboicaGermany: wandelroeschenGuinea: boulé kognokogno portoHaiti: bonbonierherbe au diableherbe bourriqueherbe à plombHawaii: lakanaIndia: bandsnagaairiphullakiputustantbiIndonesia: boenga pagarboenga pagarchentekembang satikkembang telekoblo

puchenganpuyengansaliarasaliyeresliyaratahi agamtai hayamtai kotoktelekantembelektembelekanteterapanwaungwileranKiribati: te kaibuakaMadagascar: fankatavinakohofotatramandadriekoradredrekarajejekaramityMalaysia: bunga asam senyurbunga pagarbunga tahi anjingbunga tahi asubunga tahi ayam busok

Binggeli 2005 Crop Protection Compendium - Lantana camara L. 1

bunga tahi ayamtahi ayam munaiMauritius: vieille filleNicaragua: cuasquitoPhilippines: bahug-bahugsapinitPuerto Rico: cariaquilloRyukyu Archipelago: shichi-hengeSaint Helena: wild currantSamoa:

LantanalatanaSouth Africa: boesmandruiwecherry-piecommon lantanagewone lantanagomdaggasumbavoelbrandewynwild lantanawilderoosmarynyellow sageSri Lanka: ganda-panagarda-pana

genda-panakatu-hingururata-guruton-kinnaThailand: kamkungpaka krawngpha-ka-krongTonga: TalamoatalatalaVietnam: thom oiZimbabwe: 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 hostsAnanas 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 descriptorsPrincipal weed in: managed forests; natural forests; wastelands


GEOGRAPHIC DISTRIBUTION

Notes on distributionL. 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 introduced (1692) Burkill, 1935

Italy present introduced Tutin et al., 1972 [Portugal]

Azores localized introduced Tutin et al., 1972 Madeira localized introduced invasive Press & Short, 1994

Spain localized introduced not invasive Sobrino et al., 2002

Asia Bangladesh present introduced invasive Islam et al., 2001

Brunei Darussalam present introduced invasive Holm et al., 1979; Waterhouse, 1993

Cambodia present introduced invasive Waterhouse, 1993 China present introduced invasive Holm et al., 1979; Corlett, 1992

Hong Kong present, few occurrences

introduced (ca 1851) invasive Holm et al., 1979

Taiwan widespread introduced invasive Xie Yan et al., 2001

India present introduced (1809) invasive Burkill, 1935

Andhra Pradesh present introduced invasive Rawat, 1997 Assam widespread introduced invasive Gujral & Vasudevan, 1983 Bihar widespread introduced invasive Gujral & Vasudevan, 1983 Delhi widespread introduced invasive Gujral & Vasudevan, 1983 Himachal Pradesh widespread introduced invasive Gujral & Vasudevan, 1983 Indian Punjab widespread introduced invasive Gujral & Vasudevan, 1983 Jammu and Kashmir widespread introduced invasive Gujral & Vasudevan, 1983 Karnataka widespread introduced invasive Gujral & Vasudevan, 1983 Madhya Pradesh widespread introduced invasive Gujral & Vasudevan, 1983 Maharashtra present introduced invasive Sinha & Sharma, 1984 Orissa present introduced invasive Sinha & Sharma, 1984 Rajasthan widespread introduced invasive Gujral & Vasudevan, 1983 Tamil Nadu widespread introduced invasive Nair & Henry, 1983 Uttar Pradesh widespread introduced invasive Gujral & Vasudevan, 1983


West Bengal present introduced invasive Sinha & Sharma, 1984 Indonesia present introduced invasive Waterhouse, 1993

Java widespread introduced invasive Smiet, 1992 Kalimantan widespread introduced invasive Holm et al., 1979 Sulawesi widespread introduced invasive Whitten et al., 2002

Israel present, few occurrences introduced Holm et al., 1979

[Japan] Ryukyu Archipelago localized introduced invasive Walker, 1976

Malaysia localized introduced invasive Holm et al., 1979; Waterhouse, 1993

Myanmar present introduced invasive Waterhouse, 1993

Philippines widespread introduced (ca 1840) invasive Burkill, 1935; Holm et al., 1979;

Waterhouse, 1993 Saudi Arabia present introduced Day et al., 2003

Singapore present introduced invasive Waterhouse, 1993; Baretto et al., 1995

Sri Lanka widespread introduced (ca 1826) invasive Morton, 1994; Evans, 1999

Thailand present introduced invasive Holm et al., 1979; Waterhouse, 1993

Timor-Leste widespread introduced invasive McWilliam, 2000 Turkey widespread introduced invasive Holm et al., 1979

Vietnam present introduced invasive Holm et al., 1979; Waterhouse, 1993

Africa Angola present introduced invasive Day et al., 2003

Cape Verde present introduced (ca 1851) invasive Chevalier, 1935

Comoros widespread introduced invasive Roby & Dossar, 2000 Congo Democratic Republic present introduced invasive Day et al., 2003

Côte d'Ivoire present introduced invasive Holm et al., 1979

Ethiopia widespread introduced invasive Binggeli & Desalegn Dessissa, 2002

Gabon present introduced invasive Barreto et al., 1995 Gambia present introduced invasive Robinson, 2001 Ghana widespread introduced invasive Holm et al., 1979 Guinea present introduced invasive Schnell, 1950 Kenya widespread introduced invasive Holm et al., 1979 Liberia present introduced invasive Holm et al., 1979

Madagascar widespread introduced invasive Holm et al., 1979; Binggeli, 2003

Mauritius widespread introduced invasive Macdonald et al., 1991 Rodriguez Island widespread introduced invasive Strahm, 1989


Mozambique widespread invasive Holm et al., 1979 Namibia present introduced invasive Bromilow, 1995 Nigeria widespread introduced invasive Holm et al., 1979 Saint Helena widespread introduced invasive Cronk, 1989 Senegal present introduced invasive Holm et al., 1979 Seychelles present introduced invasive Gerlach, 1993 South Africa widespread introduced invasive Erasmus et al., 1993 Sudan present introduced invasive Barreto et al., 1995 Swaziland present introduced invasive Robertson et al., 2001 Tanzania widespread introduced invasive Holm et al., 1979 Uganda widespread introduced invasive Holm et al., 1979 Zambia present introduced invasive Holm et al., 1979 Zimbabwe widespread introduced invasive Holm et al., 1979 Central America & Caribbean Antigua and Barbuda present native Francis et al., 1994

Barbados widespread native Gooding et al., 1965; Morton, 1994

Costa Rica widespread native Schemske, 1983 Cuba present native Sharma et al., 1988 Dominican Republic present native Barreto et al., 1995 El Salvador present native Day et al., 2003 Guatemala present native Holm et al., 1979 Haiti present native Day et al., 2003 Honduras present native Holm et al., 1979 Jamaica widespread native Adams, 1976 Nicaragua widespread native Holm et al., 1979 Panama present native Holm et al., 1979 Puerto Rico localized native Holm et al., 1979 Trinidad and Tobago widespread native Holm et al., 1979 United States Virgin Islands present native Morton, 1994

North America [USA]

Alabama present introduced Doren et al., 2002 California present introduced Morton, 1994 Georgia (USA) present introduced Doren et al., 2002

Hawaii widespread introduced (1859) invasive Davis et al., 1992

Louisiana present introduced Doren et al., 2002 Mississippi present introduced Doren et al., 2002 North Carolina present introduced Morton, 1994 Oklahoma present introduced Doren et al., 2002 South Carolina present introduced Doren et al., 2002


Texas present introduced Morton, 1994 Utah present introduced Doren et al., 2002

South America Argentina present native Morton, 1994 Bolivia present native Holm et al., 1979 [Brazil]

Alagoas present native Lorenzi, 1983 Amazonas present native Barreto et al., 1995 Bahia present native Barreto et al., 1995 Ceara present native Lorenzi, 1983 Espirito Santo present native Lorenzi, 1983 Goias present native Lorenzi, 1983 Maranhao present native Lorenzi, 1983 Matto Grosso do Sul present native Lorenzo, 1983 Matto Grosso present native Sharma et al., 1988 Minas Gerais present native Lorenzi, 1983 Paraiba present native Lorenzi, 1983 Parana present native Lorenzi, 1983 Pará present native Lorenzi, 1983 Pernambuco present native Lorenzi, 1983 Piauí present native Lorenzi, 1983 Rio Grande do Norte present native Lorenzi, 1983 Rio Grande do Sul present native Lorenzi, 1983 Rio de Janeiro present native Barreto et al., 1995 Santa Catarina present native Lorenzi, 1983 Sao Paulo present native Lorenzi, 1983 Sergipe present native Lorenzi, 1983

Colombia present native Morton, 1994 Ecuador present native Barreto et al., 1995

Galapagos Islands widespread introduced invasive Cruz et al., 1986 Peru present native Morton, 1994 Venezuela present native Morton, 1994 Oceania American Samoa widespread introduced invasive Thaman, 1974

Australia widespread introduced (1841) invasive Swarbrick, 1986

Australian Northern Territory localized introduced invasive Swarbrick, 1986

New South Wales widespread introduced invasive Swarbrick, 1986 Queensland widespread introduced invasive Swarbrick, 1986

South Australia localized introduced not invasive Swarbrick, 1986

Victoria localized introduced not invasive Swarbrick, 1986


Western Australia localized introduced invasive Swarbrick, 1986 Belau widespread introduced invasive Meyer, 2000 Cook Islands widespread introduced invasive Thaman, 1974 Federated states of Micronesia widespread introduced invasive Meyer, 2000

Caroline Islands widespread introduced invasive Thaman, 1974 Fiji widespread introduced invasive Mune & Parham, 1967

French Polynesia widespread introduced (1843) invasive Meyer, 2000

Guam localized introduced invasive Thaman, 1974 Nauru widespread introduced invasive Meyer, 2000

New Caledonia widespread introduced (ca 1883) invasive Heckel, 1911

New Zealand widespread introduced invasive Holm et al., 1979 Niue widespread introduced invasive Meyer, 2000 Norfolk Island widespread introduced invasive Swarbrick, 1986 Northern Mariana Islands widespread introduced invasive Meyer, 2000

Papua New Guinea widespread introduced invasive Holm et al., 1979 Pitcairn Islands widespread introduced invasive Florence et al., 1995 Samoa widespread introduced invasive Thaman, 1974 Solomon Islands present introduced invasive Swarbrick, 1986 Tonga widespread introduced invasive Thaman, 1974 Vanuatu widespread introduced invasive 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


(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.


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


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 specialistNatural 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 AsiaCharidotis pygmaea Leaves Coelocephalapion Inflorescence, Leaves Cremastobombycia lantanella Leaves Diastema tigris Leaves Ectaga garcia Leaves Crocidosema lantana MicronesiaEutreta xanthochaeta (gall fly, lantana) Fruits/pods HawaiiGeraeus nr. curvispinis Inflorescence Hepialus Stems Hypena laceratalis Leaves Langsdorfia franckii Roots Lantanophaga pusillodactyla Inflorescence MicronesiaLeptobyrsa decora Leaves Longitarsus Leaves Neogalea sunia (lantana stick, caterpillar (USA))

Leaves

Octotoma championi Leaves Octotoma pliculata Leaves Octotoma scabripennis (leaf beetle, lantana) Leaves New Caledonia; South AfricaOmophoita albicollis Inflorescence, Leaves Ophiomyia camarae Leaves Ophiomyia lantanae (fly, lantana seed) Fruits/pods,

Inflorescence Asia; Micronesia; South Africa

Plagiohammus spinipennis Stems Pseudopyrausta acutangularis Leaves


http://LaunchNENbookmark/

Salbia haemorrhoidalis Leaves South AfricaStrymon bazochii Fruits/pods,

Inflorescence Teleonemia elata Leaves Teleonemia harleyi Leaves Teleonemia prolixa Leaves Teleonemia scrupulosa (lantana, bug) Leaves Micronesia; South AfricaTeleonemia validicornis Leaves Thecla Leaves Strymon bazochii Leaves Uroplata girardi (lantana, hispid) Leaves Micronesia; New Caledonia; South

AfricaUroplata lantanae Leaves

Additional natural enemies (source - data mining)Natural enemy Pest stage attacked Biological control in:

Herbivores:Hypena strigatus (lantana defoliator, caterpillar)

South Africa

Platyptilia pussilidactyla

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.


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 human-dominated 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).


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


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 well-known 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


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


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 fruit-feeding 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


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)

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