Lantana camara
(lantana)

Pictures

Picture	Title	Caption	Copyright
Title Flowers Caption Lantana camara (lantana); flowers. Copyright ©CABI/Arne Witt	Flowers	Lantana camara (lantana); flowers.	©CABI/Arne Witt
Title Flowers Caption Lantana camara (lantana); flowers. Copyright ©CABI/Arne Witt	Flowers	Lantana camara (lantana); flowers.	©CABI/Arne Witt
Title Habit Caption Lantana camara (lantana); habit, showing flowers and foliage, as well as the thorny stems. Copyright ©CABI/Arne Witt	Habit	Lantana camara (lantana); habit, showing flowers and foliage, as well as the thorny stems.	©CABI/Arne Witt
Title Invasive habit Caption Lantana camara (lantana); invasive habit. Ethiopia. July 2014. Copyright ©CABI/Arne Witt	Invasive habit	Lantana camara (lantana); invasive habit. Ethiopia. July 2014.	©CABI/Arne Witt
Title Invasive habit. Caption Lantana camara (lantana); invasive habit, on the edge of the Queen Elizabeth National Park, Uganda. January 2005. Copyright ©CABI/Arne Witt	Invasive habit.	Lantana camara (lantana); invasive habit, on the edge of the Queen Elizabeth National Park, Uganda. January 2005.	©CABI/Arne Witt
Title Leaf Caption Lantana camara (lantana); leaf, backlit, showing venation. Copyright ©CABI/Arne Witt	Leaf	Lantana camara (lantana); leaf, backlit, showing venation.	©CABI/Arne Witt
Title Fruits Caption Lantana camara (lantana); fruits, unopened inflorescence and leaves. Copyright ©CABI/Arne Witt	Fruits	Lantana camara (lantana); fruits, unopened inflorescence and leaves.	©CABI/Arne Witt

Identity

Preferred Scientific Name

• Lantana camara L.

Preferred Common Name

• lantana

Other Scientific Names

• Camara vulgaris Benth.
• Lantana antidotalis Thonning (1827)
• Lantana camara var. aculeata
• Lantana crocea Jacq.
• Lantana glandulosissima Hayek
• Lantana mexicana Turner
• Lantana mixta Medik.
• Lantana moritziana Otto & A.Dietr.
• Lantana sanguinea Medik.
• Lantana scabrida Ait.
• Lantana spinosa L. ex Le Cointe
• Lantana undulata Raf.
• Lantana urticifolia Mill.
• Lantana x aculeata f. crocea (Jacq.) Voss

International Common Names

• English: arch man; common lantana; large leaf lantana; pink-flowered lantana; prickly lantana; red sage; red-flowered sage; shrub verbena; tickberry; white sage; wild sage; yellow sage
• Spanish: camar; cariaquillo; cinco cincos; cinco negritos; comida de paloma; corroncho; cuasquito; filigrana; frutilla; jaral; jarrila; mora de caballo; morita; palo del diablo; santo negrito; soterre; venturosa
• French: corbeille d’or; galabert; lantanier; mille fleurs; vieille fille
• Chinese: ma ying dan

Local Common Names

• Brazil: camara; cambara de espinho
• Cambodia: ach mann
• Cook Islands: ranatana; tataramoa
• Costa Rica: cinco negritos; flor de duende; tres colores
• El Salvador: bandera española
• Fiji: kauboica
• French Polynesia: tatara moa
• Germany: wandelroeschen
• Guinea: boulé kogno; kogno porto
• Haiti: bonbonier; herbe à plomb; herbe au diable; herbe bourrique
• India: bands; nagaairi; phullaki; putus; tantbi
• India/Assam: guphul
• Indonesia: 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
• Japan: shichihenge
• Japan/Ryukyu Archipelago: shichi-henge
• Kiribati: te kaibuaka
• Lesser Antilles: measle bush; rangoat leaf; sauge; scrubby tree
• Madagascar: fankatavinakoho; fotatra; mandadrieko; radredreka; rajejeka; ramity
• Malaysia: bunga asam senyur; bunga pagar; bunga tahi anjing; bunga tahi asu; bunga tahi ayam; bunga tahi ayam busok; tahi ayam munai
• Mauritius: vieille fille
• Mexico: alantana; alfombrilla hedionda (Michoacán); carrasposa; confite; confituria; confiturilla (Sonora-Chihuahua); confiturio (Baja California); flor de San Cayetano; lampana; matizadilla; pasaruin; scrubby cap; sonora roja (Sinaloa); uña de gato (Morelia)
• Micronesia, Federated states of: randana (Pohnpei)
• Nicaragua: cuasquito
• Philippines: bahug-bahug; sapinit
• Portugal: cambará
• Puerto Rico: cariaquillo
• Saint Helena: wild currant
• Samoa: Lantana; latana
• South Africa: boesmandruiwe; cherry-pie; common lantana; gewone lantana; gomdagga; sumba; voelbrandewyn; wild lantana; wilderoosmaryn; yellow sage
• Spain: bandera; banderita; espuela de galán
• Sri Lanka: ganda-pana; garda-pana; genda-pana; katu-hinguru; rata-guru; ton-kinna
• Thailand: kamkung; paka krawng; pha-ka-krong
• Tonga: Talamoa; talatala
• USA/Hawaii: lakana; lanakana; mikinolia hihiu; mikinolia kuku
• Venezuela: cariaquillo; cariaquito
• Vietnam: thom oi
• Zimbabwe: chiponiwe

EPPO code

• LANCA (Lantana 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 as a hedge plant and has since been reported as extremely weedy and invasive in many countries. It is generally deleterious to biodiversity and has been reported as an agricultural weed resulting in large economic losses in a number of countries. In addition to this, it increases the risk of fire, is poisonous to livestock and is a host for numerous pests and diseases. L. camara is difficult to control. In Australia, India and South Africa aggressive measures to eradicate L. camara over the last two centuries have been largely unsuccessful, and the invasion trajectory has continued upwards despite control measures. This species has been the target of biological control programmes for over a century, with successful control only being reported in a few instances. 

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Lamiales
•                         Family: Verbenaceae
•                             Genus: Lantana
•                                 Species: Lantana camara

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 L. 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 (Stirton, 1979; Gujral and Vasudevan, 1983; Scott et al., 1997). Two groups are often recognised: one with few or no spines commonly found in the neotropics and one with spines in other parts of the world where the species is troublesome (Howard, 1970; Swarbrick, 1986). In the Pacific Islands the most common variety is the prickly L. camara var. aculeata (Thaman, 1974).

Description

L. camara is a medium-sized perennial aromatic shrub, 2-5 m tall, with quadrangular stems, sometimes having prickles. The posture may be sub-erect, scrambling, or occasionally clambering (ascending into shrubs or low trees, clinging to points of contact by means of prickles, branches, and leaves). Frequently, multiple stems arise from ground level. The leaves are generally oval or broadly lance-shaped, 2-12 cm in length, and 2-6 cm broad, having a rough surface and a yellow-green to green colour. The flat-topped inflorescence may be yellow, orange, white, pale violet, pink, or red. Flowers are small, multicoloured, in stalked, dense, flat-topped clusters to 4 cm across. Fruit is a round, fleshy, 2 seeded drupe, about 5 mm wide, green turning purple then blue-black (similar in appearance to a blackberry).

Plant Type

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). This species has been widely promoted as an ornamental since the early 1800s and it is widely naturalized throughout the Neotropics. It is present on all continents expect Antartica. It has become very widespread in Australia, India and South Africa, infesting millions of hectares of land (Bhagwat et al., 2012).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

L. camara has been introduced throughout the tropics and subtropics, often used as a hedge plant, and is commonly grown in the temperate zone. Although first cultivated in Europe during the late seventeenth century, reaching Calcutta in 1809 (Burkill, 1935), it was mostly introduced throughout the tropics during the later part of the nineteenth century and a number of cultivars and forms were subsequently disseminated (Howard, 1970). 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).

Risk of Introduction

The main spread of L. camara into new countries is via the horticultural trade, spreading numerous different varieties of this species. Once introduced into an area the seeds of L. camara are readily dispersed into new areas by birds. L. camara has already spread widely, but there is potential for its range to expand further under future climate change. In Australia it is found in almost all the climatically suitable habitat, but under future climate change it could expand into new areas in Victoria, South Australia and Tasmania (Taylor et al., 2012; Taylor and Kumar, 2013).

Habitat

L. camara has a wide environmental tolerance and occurs in a variety of habitats. These include wastelands, rainforest edges and beachfronts (ISSG, 2015). It also grows well in disturbed areas such as roads, railways and areas recovering from fire or logging (ISSG, 2015). This species can tolerate some shade but grows best in open unshaded regions. It cannot directly colonise intact forests but instead grows at forest edges and spreads when gaps are created (ISSG, 2015).

Habitat List

Hosts/Species Affected

L. camara is an agricultural weed that can cause dramatic losses in yields. In Australia, it was reported that L. camara infested 4 million ha of pasture (Parsons and Cuthbertson, 1992). A number of plants affected by L. camara are listed in the "Host Plants/Plants Affected" table below.

Host Plants and Other Plants Affected

Biology and Ecology

Genetics

The known chromosome numbers of L. camara are 2n = 22, 33, 44, 55, but most invasive varieties 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 two months of the rainy season.

In the highlands of western Kenya an investigation of leaf decomposition found that after seven 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 12-34% 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. Projections of seed survival indicate that L. camara seeds could survive for up to 11 years under natural rainfall conditions in Australia (Vivian-Smith and Panetta, 2009).

In addition to spread 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 1,400 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. Studies undertaken by Muvengwi and Ndagurwa (2015) on soil seed bank dynamics and soil nutrient concentrations in the wetlands of New Gada in Zimbabwe, suggest that L. camara-invaded soil had significantly higher levels of calcium, magnesium, sodium and ammonium and lower levels nitrate than non-invaded soils. 

L. camara has a marked ability to compensate for herbivory as plants survived experimental defoliation for two 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.

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free

Soil reaction

• acid
• neutral

Soil texture

• heavy
• medium

Special soil tolerances

• infertile

Natural enemies

Notes on Natural Enemies

The alkaloid-rich leaves of L. camara make it 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. 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). In Mexico a stem sap-sucking membracid bug, Aconophora compressa, causes considerable dieback of stems (Swarbrick et al. 1995).

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), Breeÿen et al. (2000), Thomas and Ellison (2000), Trujillo and Norman (1995).

Means of Movement and Dispersal

Natural Dispersal

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). In the Kruger National Park, South Africa, L. camara has primarily spread along rivers (Vardien et al. 2012).

Vector Transmission

The seeds of L. camara 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, Acridotheres tristis (Atkinson and Atkinson, 2000). In the Galapagos Islands it is one of the most dispersed alien plants, being mainly dispersed by two lizard species, and to a minor extent by the birds Myiarchus magnirostris and Mimus melanotis (Heleno et al., 2013). There are also reports of seeds being dispersed by sheep and goats.

Accidental Introduction

Accidental introduction of L. camara 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.

Pathway Causes

Pathway Vectors

Impact Summary

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). In Fiji it is a major weed of coconut plantations, pastures, neglected arable land and waste places (Mune and Parham, 1967). 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 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 Kenya, L. camara is poisonous to livestock and also a habitat for tsetse flies (IPPC-Secretariat, 2005). 

Environmental Impact

Impact on Habitat

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 (Heckel, 1911). Its extensive seed production favours rat populations.

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 5-6 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. There are many unsubstantiated statements suggesting that L. camara slows erosion (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 may have some potential in forest restoration (Burkill, 1935).

Impact on Biodiversity

L. camara can readily hybridise 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; Gooden et al., 2009) but there are some occasional references to regeneration of some tree species under its canopy (e.g. Burkill, 1935). Turner and Downey (2010) used L. camara as a case study for their methodology to identify the native biodiversity threatened by an invasive plant. They identified 275 and native plants and 24 native animals requiring protection from L. camara invasions in Australia. 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 thickets of L. camara 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).

Threatened Species

Social Impact

Stands of L. camara 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).

Risk and Impact Factors

• Proved invasive outside its native range
• Highly adaptable to different environments
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Highly mobile locally
• Has high reproductive potential

• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Modification of fire regime
• Negatively impacts agriculture
• Negatively impacts forestry
• Negatively impacts human health
• Negatively impacts animal health
• Reduced native biodiversity
• Threat to/ loss of native species

• Allelopathic
• Competition - monopolizing resources
• Competition - shading
• Competition - smothering
• Competition
• Pest and disease transmission
• Hybridization
• Produces spines, thorns or burrs

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately
• Difficult to identify/detect as a commodity contaminant
• Difficult/costly to control

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 fence of L. camara to protect crops from domestic animals was taken up by local villagers in the 1990s but this quickly led to the loss of rough grazing land through the rapid spread of this species (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 straw from L. camara mixed with dung for biogas production and using the twigs as fuel. There is some evidence, although conflicting in nature, that extracts from L. camara 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, Solanum tuberosum, with leaves of L. camara almost eliminates damage by the potato tuber moth Phthorimaea operculella (Lal, 1987).

Uses List

Animal feed, fodder, forage

• Fodder/animal feed

Environmental

• Boundary, barrier or support
• Erosion control or dune stabilization

Fuels

• Biofuels
• Fuelwood

General

• Ornamental

Human food and beverage

• Honey/honey flora

Materials

• Pesticide
• Poisonous to mammals

Medicinal, pharmaceutical

• Traditional/folklore

Detection and Inspection

L. camara is conspicuous due to its attractive and multicoloured floral displays and is a well-known species throughout the tropics.

Similarities to Other Species/Conditions

Although large stands of weedy varieties of L. camara are easily recognised, it is in fact a variable polyploid complex of interbreeding taxa resulting from hybridisation 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, L. depressa, with which it has extensively hybridised (Langeland and Burks, 2000).

Prevention and Control

Cultural Control

Being poisonous to livestock 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).

Osunkoya et al. (2013) suggest from studies and simulation models in Queensland, Australia that periodic burning could control the weed in forests within 4-10 years if fire frequency is at least every two years. On farms, site-specific control may be achieved by 15 years if the biennial fire frequency is tempered with increased burning intensity.

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 colonised via seeds originating from distant parents or from sprouting roots. Dohn et al. (2013) recommend hand pulling for creating firebreaks or where minimizing damage to native species is paramount.

Chemical Control

The Australian experience in controlling L. camara, reviewed by Swarbrick et al. (1995), indicates that some herbicides are more effective on particular forms of L. camara. The most effective herbicides belong to the phenoxy acid (2,4-D, dichloroprop and MCPA), benzoic acid (dicamba) and pyridine groups. Glyphosate, 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 been reported by Hannan-Jones (1998).

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, Himachal Pradesh, was effective and economical using glyphosate sprayed on to regenerated growth, cut four months previously (Rana and Singh, 1999).

In Queensland, Dohn et al. (2013) suggest that foliar spraying with a glyphosate-based herbicide is the most efficient treatment for combating large infestations of L. camara. In Florida, Ferrell et al. (2011) report that this species can be effectively controlled by two applications of fluroxypyr, two applications of fluroxypyr+aminopyralid, or a single application of aminocyclopyrachlor. 

L. camara is resistant to triclopyr, a widely used herbicide for woody weed control (Goodall and Naude, 1998).

Biological Control

Worldwide, well over 200 releases of biocontrol agents 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. 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. 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; Denton et al., 1991; Davis et al., 1992; 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 to the control of L. camara 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 U. 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 L. camara 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 the insect species varied between islands and between varieties of L. camara. Greater success appears to have been achieved in drier areas.

In Uganda, the introduction of T. 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 L. camara had been controlled. Subsequently other agents for L. camara 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).

Recent releases of arthropod biocontrol agents in Australia (Queensland and New South Wales) to control L. camara include the treehopper Aconophora compressa (first in 1995), the mirid Falconia intermedia (during 2000-2004), and the leaf miner Ophiomyia camarae (first in 2007). Weather conditions and other factors have resulted in poor establishment or low levels of damage so far (Taylor et al., 2008).

Hersula and Hill (2012) report that populations of F. intermedia released in South Africa to control L. camara had disappeared after initially building up to high densities. It is suggested that some L. camara varieties possess factors enabling them to resist feeding activities after the initial attack. Biological control of L. camara in South Africa is reviewed by Moran et al. (2011), who suggest that it plays a subsidiary role in support of mechanical and chemical control, but that cost benefits justify the continued development of new agents.

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 U. 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. Julien and Griffiths (1998) showed that different cultivars display differences in susceptibility to insect herbivores.

A potential pathogen of L. camara (spreading in Hawaii) was identified by Trujillo and Norman (1995) as a leaf-spot fungus, Septoria sp., from Ecuador. Various other pathogens with apparently excellent potential to control a wide range of cultivars have been identified by Barreto et al. (1995) and Thomas and Ellison (2000). In South Africa, permission was granted in 2001 to release the fungus Mycovellosiella lantanae var. lantanae, collected from Florida, USA (Breeÿen et al., 2000, Breeÿen, 2004). In 2001 the rust fungus Prospodium tuberculatum (ex Brazil) was the first pathogen to be released in Australia for biological control of L. camara (Tomley and Riding, 2002; Ellison et al., 2006; Thomas et al., 2006). Drought conditions affected its establishment and spread from the release sites in Queensland and New South Wales and incidence is generally low, although some leaf drop has been observed (Taylor et al., 2008).

In May 2015, the first release of P. lantanae was made on New Zealand’s North Island. The rust had previously been screened by CABI scientists in the UK for its host specificity before being transferred to Landcare’s Plant Pathogen Containment Facility in Auckland. The blister rust is able infect leaves, petioles and stems and can cause systemic infections that lead to stem dieback and defoliation.

In addition, a second rust, P. tuberculatum (the same isolate released in Australia in 2001) was released in New Zealand as part of the same initiative. The leaf rust pathogen causes leaf death and defoliation and as it is subtropical, it is expected to be less dependent on high humidity to compliment P. lantanae in different climatic conditions (Anon, 2015).

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Contributors

07/12/15 Updated by:

Sarah E. Thomas, CABI, UK

23/09/13 Updated by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps