Protection Quarterly Vol.10(2) 1995 82 In weedy forms about half of the flow- The Biology of Australian Weeds ers usually produce a single-seeded fleshy spherical 5–7 mm in diameter, 25. camara L. at first hard and green but ripening to purple or black and then consisting of a J.T. SwarbrickA, B.W. WillsonB and M.A. Hannan-JonesB. thin skin containing a purple pulp around A Weed Science Consultancy, 15 Katoomba Crescent, Toowoomba, the stony, pear-shaped, 3–4 mm diameter Queensland 4350, . . The non-weedy forms set very few . Each seed contains one or some- B Alan Fletcher Research Station, PO Box 36, Sherwood, Queensland 4075, times two embryos, and in the latter case Australia. both may germinate. The root system typically consists of a short tap root with laterals, which divide Name repeatedly to form a root mat. Lantana was known by at least tall but capable of becoming a liane up to does not sucker from damaged or broken five polynomial descriptive names, start- 15 m tall if given support (Figure 1). roots, but will regrow vigorously from the ing with Lantana, Viburnum and The arching, scrambling or prostrate base of the stem and more slowly from Periclymenum, before Linnaeus gave it its stems are initially 4-angled but become rooted horizontal stems in contact with binomial name in 1753. He retained cylindrical and up to 15 cm thick with moist soil. Lantana (origin obscure) and described a age. Young stems are hairy and in the number of species including camara (a weedy forms carry sharp recurved prick- Variation within the species West Indian name) and aculeata (prickly), les along the angles, whilst those of the Lantana is an aggregate species, derived a species now included within the non-weedy forms are rounder, more slen- through horticultural and natural hy- L. camara complex (L. camara sensu lato). der and without prickles. bridization, selection and somatic muta- Linnaeus’ L. camara was non-prickly The opposite pairs of ovate leaves oc- tion from a number of similar and prob- whilst his L. aculeata was prickly; both cur on 1–3 cm long petioles. Each leaf is ably closely related (though spatially were described from European garden 3–10 cm long and 2–6 cm wide, with a separated) tropical American species. material (Schauer 1851). bluntly pointed, rounded or heart-shaped This has resulted in many hundreds of Lantana has become the common name base, finely toothed margin and bluntly forms, which are also referred to as for the most abundant and complex spe- pointed tip. The leaves are rough and cultivars and varieties (Howard 1969). cies within the genus. It contains several finely hairy on both surfaces and strongly The forms vary in ploidy, bush shape, hundred wild and cultivated forms. veined below, and have a strong charac- colour, prickliness, response to the teristic smell when crushed. The upper environment and natural enemies, and Description epidermis carries very few stomata. Both chemical composition, including toxicity Lantana is a brittle, much branched, the stomata and the trichomes from both to . At least 29 forms have become thicket-forming shrub, normally 2–4 m surfaces of leaves have been related to tax- naturalized in eastern Australia (Smith onomy (Raghavan and Arora and Smith 1982) and probably others in 1960, Shah and Mathew 1982). the Northern Territory and Western Aus- Inflorescences develop in the tralia. A number of non-weedy forms are axils of young leaves. They are sold by nurseries. Smith and Smith (1982) compact, flat to dome-shaped considered that 19 forms were sufficiently and 2–3 cm across and each is common in eastern Australia to be eco- supported by a finely hairy nomically important as either weeds or 2–8 cm long peduncle with a poisonous . club-shaped tip. Each inflores- The basic chromosome number of cence contains 20–40 sessile Lantana is 11, with 12 being the basic , each subtended by a number for the rest of the genus stiff deciduous bract. The (Raghavan and Arora 1960). Diploid, flower buds are angular and triploid, tetraploid, pentaploid and tightly packed, and open from hexaploid forms of lantana have been re- the outside of the inflorescence ported in (Stirton 1977, Spies towards the centre. Each and Stirton 1982, Spies 1984), and triploid, flower has a 9–12 mm long tetraploid and pentaploid in In- tube and four unequal rounded dia (Sen and Sahni 1955). Raghavan and 4–8 mm across. Newly Arora (1962) showed that Indian diploids opened flowers normally have were small, stunted and very rare and yellow throats, and the petals that the weedy forms were mostly vary from white to cream, yel- tetraploids with some triploids. Koshoo low, pink, orange, red or pur- and Mahal (1967) claimed that the breed- ple. After flowers ing system combines sexual, semisexual darken, lose the yellow centre and asexual (apomictic) systems, and and fall, except in non-weedy Stirton (1977) stated that polyploidy has forms which mostly fail to set developed through both auto- and allo- seed and retain their flowers polyploidy. longer. Pollen grains are al- Most of the weedy forms of lantana in most globular, tricolpate and South Africa and and probably Aus- 30–35 µm across and are finely tralia appear to be tetraploids, whilst most Figure 1. Lantana (Lantana camara L.) (Line textured (d’Almeida and of the non-weedy ornamental forms ap- drawing by Dr. G. Scott). Roland-Heydacker 1985). pear to be triploids. 83 Plant Protection Quarterly Vol.10(2) 1995 After growing many weedy forms of larger bushes, longer internodes and leaf Adelaide (Bailey 1841), and was followed lantana together in South Africa to study blades, prickly stems and high levels of by many others. The plant was quickly their cytology and the morphology of fruit production, whereas most of the or- spread northwards along the coast, with their inflorescences, bracts and flowers, namental forms produced smaller the first Australian collections of appar- Spies (1984) concluded that ‘plants on dif- bushes, had shorter internodes and ently naturalized lantanas from the Bris- ferent ploidy levels are morphologically leaves, lacked prickles, and while produc- bane River in 1861 and near in similar’. A similar study of Indian plants ing similar numbers of flowers, set very 1862 (Swarbrick 1985). By 1897 lantana by Natarajan and Ahuja (1957) showed few fruit. The ornamental form ‘Drap d’or’ was described as being ‘a huge rambling that non-prickly forms occurred at 2n, 3n occupied an anomalous position since it shrub and a most troublesome weed and 4n levels, flower colour varied within showed the large bush size, long which has spread to an alarming extent, all ploidy levels, and all ploidy levels con- internodes and large leaves of the weedy and forms impenetrable thickets on the tained forms with both nil and good seed group but the lack of prickles and low banks of streams, deserted farms and the set. seed production of the ornamental group, edges of scrubs’ in the Brisbane area and Tandon and Bali (1955) compared dip- suggesting significant differences in its as being ‘equally abundant about Port loid and triploid forms and found that the derivation from both other ornamental Jackson’ (Bailey 1897). Lantana appears to latter was more vigorous, had larger and and weedy forms. have been grown in Western Australia by darker leaves, produced more and larger 1875 (Swarbrick 1985) but may have only flowers and, unlike the diploid, was largely History spread to the Northern Territory during sterile. Raghavan and Arora (1960) found Lantana was probably first introduced or since the Second World War (I.L. Miller no correlations between flower colour or into about 1640 from Brazil personal communication). Further expan- lower leaf epidermal patterns and five (Howard 1969), where both prickly and sion into previously unoccupied areas is levels of ploidy. non-prickly taxa are considered to be na- probably still occurring, often assisted by Swarbrick (unpublished) compared the tive, common and widespread (Schauer land clearing or other human disturbance morphology of seven weedy forms from 1851). Introductions continued through- (Humphries and Stanton 1992). Smith’s collection at the CSIRO in Bris- out the 17th, 18th and 19th centuries bane (now destroyed) and six ornamental (Swarbrick 1985) resulting in an aggre- Geographical distribution forms from commercial nurseries under gate species of mixed parentage (Howard uniform conditions in south-eastern 1969, Stirton 1977). Lantana became a fa- Australia Queensland. He studied bush volume, vourite hothouse plant in Europe in the Lantana is widely distributed throughout average length of the third internode from late 19th century, with 397 new varietal coastal and subcoastal areas from the shoot tip, average length of fourth names being added to nurserymen’s cata- Ulladulla in New South Wales to Cape node leaves, average number of prickles logues between 1850 and 1900 (Howard Melville in Queensland, with very scat- per 5 cm of angle on the third internode 1969). Many of these forms would have tered infestations in the Northern Terri- and average number of flowers and fruits been distributed to the colonies, including tory, Western and South Australia (Figure per inflorescence. Analysis of results Australia. 2). It is also present as a weed on both showed that the weedy forms were all The first Australian record of lantana Lord Howe and Norfolk Islands very similar and were characterized by was in 1841 at the old Botanic Gardens in (Swarbrick 1985).

Outside Australia Lantana is widely grown as an ornamen- tal shrub throughout the tropics, subtrop- ics and warm temperate zones. It is es- tablished as a weed throughout the trop- ics and subtropics, from southern USA and the Mediterranean in the north to South Africa and the northern tip of New Zealand in the south (Figure 3).

Habitat

Climatic requirements Lantana grows well over a wide range of warm temperate, subtropical and tropical climates (Gujral and Vasudevan 1983). It does not grow at temperatures below 5°C and the shoots are sensitive to frost (Thaman 1974, Winder 1980). Thaman also reported that lantana grows equally well on the drier leeward and moister windward sides of high islands across the Pacific. In south-eastern Queensland it is similarly found in moist coastal areas that naturally support rainforest and drier east- ern slopes of the Dividing Range which support open eucalypt woodland, but ceases to the west of the Dividing Range where both and frost occur. The Figure 2. The Australian distribution of lantana (Parsons and Cuthbertson upper temperature limit for lantana 1992, with permission). growth is not known. Lantana grows Plant Protection Quarterly Vol.10(2) 1995 84 Eucalypt seedlings generally fail to establish under lantana (Driscoll and Quinlan 1985), probably because of their light requirement during germina- tion. Grasses and most herbs are shaded out by dense lantana thickets, leaving bare ground when the lantana is physically or chemically re- moved (Swarbrick unpub- lished). Lamb (1988) has shown that within ten years of invasion the nature of eucalypt woodland changes signifi- cantly and possibly irrevers- ibly from dominance by native trees to dominance by the ex- otic shrub. Humphries and Stanton Figure 3. The world distribution of lantana as a weed in suitable terrestrial habitats. (1992) noted that lantana is un- (Redrawn from Holm et al. 1977, Stirton, 1977 and other sources). able to invade tropical rain for- est in northern Queensland from ‘within a few yards of the beach’ severely drained (Humphries and and only persists along edges and where (Thaman 1974) to 1000 m on the Great Di- Stanton 1992). the canopy is broken, a similar situation viding Range (M.J. Russell personal com- Lantana has a very low tolerance of soil to that found in dry scrubs in south- munication). salinity, which induces chlorosis eastern Queensland (Swarbrick unpub- Lantana grows best under conditions of (Thaman 1974) and is severely damaged lished). Many rainforest species germi- constant rainfall or where soil moisture is or killed by 7.5 mg boron per litre in sand nate and develop under lantana, eventu- available throughout the year. Humphries culture (Francois and Clark 1979). Winder ally growing through it and shading it and Stanton (1992) reported that in north- (1980) noted that lantana is very rare on out. Broadleaf privet (Ligustrum lucidum) ern Queensland it is mostly confined to hard, phosphate deficient soils of open also behaves in this way around areas receiving more than 1250 mm per eucalypt forests. Lamb (1988) has shown Toowoomba (Swarbrick unpublished). year and in the wet tropical areas its vig- that lantana increases the levels of organic Following initial disturbance lantana our and density decline with decreasing matter and nitrate nitrogen in the soils be- may, however, form a stable community rainfall. They noted that in the subtropics low well established clumps. The leaf lit- in such areas and resist rainforest re- it is rarely found west of the 750 mm ter and soil from beneath such clumps is establishment for several decades (Lamb isohyet. Bartholomew and Armstrong sometimes collected for use in gardens, 1991). Stocker and Mott (1981) showed (1978) suggested the 650 mm isohyet as where ‘lantana mulch’ is considered a de- that lantana may slow down or even its western limit in south-eastern Queens- sirable material for boosting plant growth. block grass invasion of disturbed rainfor- land. Low temperatures and dry soil in- est and that it can protect rainforest mar- dependently limit the growth of lantana. Plant communities gins from damage by low intensity fires Most susceptible areas are probably al- Parsons and Cuthbertson (1992) reported at least early in the dry season. ready infested, although further advances that lantana ‘occurs as a weed along In south-east Queensland lantana in- are likely around their fringes and per- roadsides, creek banks, fence lines, and vades and dominates native woodland haps along river systems in the Gulf of waste places and is a common compo- that has been cleared to increase its graz- Carpentaria, the Northern Territory and nent of weedy pastures (and) park ing capacity (Collins 1976). Clearing in- Western Australia. lands… and may become the dominant creases sunlight at ground level and de- understorey in open forests and tropical creases interspecific plant competition. Substrata tree plantations.’ It readily invades open Whilst this initially increases growth of Lantana grows best on rich organic soils and semi-open plant communities such native pasture species it also allows inva- developed under rainforests, although it as road verges, grasslands and wood- sion by lantana, with which the native is also reported to tolerate poor soils and lands, and proliferates under lightly grasses are unable to compete. almost pure sands (Winder 1980). Its shaded conditions such as mature hoop Many authors note that lantana is less roots tend to rot in waterlogged soils pine plantations and the early stages of common in undisturbed native communi- (Thaman 1974), and it does not become commercial exotic pine plantations. It ties than in those that have been disturbed established in sandy soils which dry out also forms monospecific stands along by cultivation, grazing, road works etc. in to depth during the dry season unless road sides and fence lines and in open which it often dominates the secondary there is a local source of soil moisture. grasslands. succession. Lantana grows on well drained clay soils Lantana initially invades woodlands as and on volcanic soils derived from basalts isolated plants under perching and roost- associations (Humphries and Stanton 1992), a situa- ing trees, from which foci it rapidly in- The of lantana are primarily distrib- tion parallelled by its vigorous growth on vades the rest of the area (Lamb 1988). uted by fruit-eating birds and mammals, high volcanic islands in Polynesia (Flor- Humphries and Stanton (1992) reported and the plant provides shelter and food for ence et al. 1983). In the wet tropics of lantana as a ‘serious invader of open for- many species of animals and small birds. northern Queensland lantana is mostly est communities’ in northern Queens- Whilst providing nectar for pollinating confined to the deeper, moister and better land, as it is for similar situations else- (see under Pollination below) it is drained metamorphic and granitic soils, where throughout its global range (e.g. toxic to many grazing animals (see under not occurring where soils are shallow and Marr 1962, Gujral and Vasudevan 1983). Detrimental importance below). 85 Plant Protection Quarterly Vol.10(2) 1995 Many species of native and exotic birds leaves and debris, sometimes developing butterflies, and various kinds of have been recorded feeding on lantana into vigorous connected daughter plants. bees (Dronamraju 1958, Schemske 1976, fruits in Australia (Liddy 1985, Loyn and The presence of a low branched tree or Kugler 1980) and by the movements of French 1991). R.J.S. Beeton (personal other support allows lantana to scramble smaller insects such as thrips within indi- communication) argues that lantana is upwards to a height of 10–15 m, in which vidual flowers (Mohan Ram and Mathur generally beneficial to wildlife in south- case the lower stems lose their branches 1984, Mathur and Mohan Ram 1986). eastern Queensland, providing feeding and assume a flexible liane-like form. Thrips were only found in yellow (newly- sites for seed-, leaf- and litter-feeding in- Established lantana bushes regrow vig- opened) flowers by Mathur and Mohan sects and shelter for small birds and orously from dormant basal buds after Ram (1978), entering through the throat mammals. Both Driscoll and Quinlan shoot removal and flower within a few and pushing past the anthers to feed on (1985) and Alcova (1987) surveyed birds months under favourable conditions. the stigmatic exudate. Thrips eat, carry and small mammals along lantana-in- and deposit pollen grains as they move fested and lantana-free transects in tall Perennation about within the flower, and carry pollen woodland in Brisbane. Dense lantana al- Lantana is a perennial shrub, constantly grains on their legs and abdomens as tered the incidence of some birds and ani- renewed at the base and long lived under they seek out other yellow-throated flow- mals, but the results were neither clearcut favourable conditions. Plants tend to die ers within the same inflorescence. nor repeatable. No differences were found only under stressful conditions including Ramachandra Rao (1920) described in the species of ground litter be- intense or prolonged drought or shading pollination of lantana by butterflies in In- tween lantana-infested and lantana-free by taller evergreen mesophytic plants. dia and Kugler (1980) pollination by bum- tall open woodland and open forest in ble, honey and other types of bees as well Brisbane by Traby (1986), although Phenology as by butterflies and moths in Crete. Ac- lantana-free areas generally had signifi- Parsons and Cuthbertson (1992) reported tivities included probing the flower tube cantly higher numbers of springtails and that lantana seedlings germinate through- by the ’s proboscis, the unconscious mites. out the year provided sufficient moisture gathering of pollen grains onto the pro- Exotic birds such as Indian mynahs are is present, with a peak after the first sum- boscis from the anthers halfway up the responsible for spreading lantana in Ha- mer rains. Initial growth is slow until the tube, and their transfer to the next waii and New Caledonia and are signifi- root system is established, after which it flower visited. Mathur and Mohan Ram cant intra- and inter-island distributors of increases as the season advances. They (1986) showed that butterflies seek nec- its seeds (Thaman 1974). Indian mynahs reported that the first flowers are pro- tar in both newly opened (unpollinated) are now spreading throughout parts of duced in the field during the second sum- and in older (pollinated) flowers, but that Australia in which lantana is already es- mer, although flowers can be produced they spend more time probing newly tablished. In South Africa lantana seeds six months after in the green- opened flowers. are spread by birds, rodents and monkeys house (Hannan-Jones unpublished). After pollination, lantana flowers lose (Wells and Stirton 1981), and in India the Lantana exhibits rapid shoot growth un- their yellow centres and usually darken in fruits are eaten by cattle, sheep, goats, der favourable conditions of soil and air colour. Mathur and Mohan Ram (1978) foxes and jackals and the seed dispersed humidity, temperature and light, resulting and Mohan Ram and Mathur (1984) ana- by many species of birds including spar- in year-round shoot production. It also lysed this colour change, in which the yel- rows, parrots and crows (Bisht and achieves marked drought tolerance under low carotenoids of the newly opened Bhatnagar 1979). The spread of lantana in temporarily dry conditions through defo- flower are masked by red . New Caledonia is assisted by the soil dis- liation. The yellow carotenoids remain dominant turbance caused by exotic ungulates in- Lantana flowers whenever soil moisture in unpollinated flowers. cluding deer, cattle, horses and goats is available and air humidity and tempera- (Thaman 1974). ture are high, resulting in almost continu- Seed development ous flowering and fruit set along the A considerable literature exists on the cy- Growth and development Queensland and New South Wales tology and seed development of lantana. coasts. A distinct flush of flowers a few Padmanabhan described the develop- Morphology weeks after soaking rains is more com- ment of the endosperm in lantana in 1959 Lantana is very plastic in its responses to mon towards the drier inland margins of and Koshoo and Mahal (1967) investi- light intensity and support. Seedling den- its Australian distribution, usually fol- gated the cytology of lantana seeds, sities vary from less than one to several lowed by significant fruit set. studying pollen viability and seed produc- per square metre. At densities of less than tion in diploids, triploids, tetraploids and 1 m-2 seedlings commonly branch near Reproduction pentaploids. All ploidy levels exhibited the base to produce rounded young some level of pollen viability and pro- plants, whilst at higher densities competi- Floral biology duced seed from open pollination. These tion for light results in more upright Parsons and Cuthbertson (1992) reported authors suggested that the seeds resulted plants. Over a number of years some that under good conditions lantana plants from normal sexual union in the diploids plants dominate and shade out the rest, flower from early summer until March or and from either facultative or obligate with each dominant plant eventually oc- April of the year after germination and apomixis at other ploidy levels, and con- cupying several square metres. Adjacent every summer thereafter from September cluded that lantana’s ‘reproductive versa- plants interlock and grow through and or October until March or April. Many au- tility’ is based on ‘a complicated mixture over each other, producing an impenetra- thors (e.g. Auld 1969) have shown, how- of sexual, semisexual and totally ble thicket several metres high with a ever, that lantana can flower throughout apomictic biotypes’. Each biotype or form canopy of active shoots. New shoots con- the year under moist, warm and well lit was thought to be the basis for the evolu- stantly arise from the base of large healthy conditions. tion of new forms, leading to the ‘intricate plants, whilst older shaded stems lose Freshly opened flowers produce nectar reticulation of taxonomic differences’ vigour and die. Seedlings germinating (Kugler 1980), which attracts many spe- within the aggregate species. under such clumps usually fail to estab- cies of insect . Both self and Stirton’s 1977 review of the literature on lish. Prostrate stems root at the nodes if cross pollination are brought about lantana threw doubt on Koshoo and they become covered by moist fallen through visits of large insects such as Mahal’s 1967 suggestion that apomixis Plant Protection Quarterly Vol.10(2) 1995 86 plays some part in seed production since germination from 29 to 47% after 85 days. productivity of pasture land. Lantana this finding was not supported by any The lantana seed also affected the ger- readily invades uncultivated pastures, other author. Spies and Stirton (1982a) mination percentage of infested fruits (see and unless controlled spreads to exclude examined embryo sac development in 20 under Response to natural enemies be- useful native grasses and fodders. The South African forms of lantana, specifi- low). plant is unpalatable and scarcely eaten, cally looking for evidence of apomixis but The influence of the fruit pulp as an in- and the trampling and deposition of dung failing to find any. Ovules from diploid, hibitor for lantana seeds does not appear by large hoofed animals are instrumental triploid and tetraploid forms all contained to have been specifically tested. Hiet in the germination and establishment of normal sexual embryos, but no normal (1944, 1946) reported significant in- new seedlings. embryo sacs were found in either creases in the germination of dry-stored Lantana contains a number of chemi- pentaploid or hexaploid forms. Forty two lantana seeds after removal of the pulp. cals toxic to animals, especially the and a half per cent of ovules contained Hannan-Jones (unpublished) observed triterpene lantadenes A and B (Everist two sexual embryo sacs, suggesting the that 44% of seeds collected from the fae- 1981). Whilst ungulates are primarily af- possibility of two embryos developing to ces of handled wild birds germinated, fected, there have been some cases of maturity within such seeds. compared with 10% for seeds from the human poisoning. All forms of lantana are Spies and Stirton (1982b) concluded same bushes which retained their pulp thought to be toxic, although the toxicity from further studies of the cytology of and 46% for similar but cleaned seeds. It varies considerably between forms South African lantanas that there are appears that the pulp around lantana (Seawright 1965). Lantana poisoning oc- large genomic differences between forms seeds serves not only to attract seed vec- curs most frequently in cattle and less of- at the same ploidy levels, indicating either tors but also to delay or even inhibit the ten in sheep, probably because few sheep the hybridization of different cultivars be- germination of seed falling under the par- are kept in areas prone to lantana inva- fore their introduction into South Africa ent bush. sion. Although photosensitization is the or a high rate of post introduction evolu- commonest symptom of lantana poison- tion within the species, or possibly both. ing in sheep, acute damage is mainly to Differences in chromosomal behaviour Parsons and Cuthbertson (1992) reported the liver and chronic damage to the liver, between the South African and Indian that new lantana stems may arise from kidneys and gut (Seawright and Hrdlicka forms indicate the introduction of differ- lateral roots in early spring and that the 1977). ent groups within the complex into the size of lantana clumps increases by Nearly 4 million ha of pasture are different countries, and it would be safe suckering. The growth of stems from thought to be infested with lantana in Aus- to assume that the Australian lantanas lantana roots has not been observed by tralia (Parsons and Cuthbertson 1992), will be different again. any of us despite close association with mostly in coastal and subcoastal Queens- lantana and careful observation over land and New South Wales. Annual Seed production and dispersal many years. Lantana bushes often have losses due to lantana in pastures are esti- The average percentage fruit set in a prostrate stems with adventitious roots mated to be A$7.7m, comprising 1500 cat- number of open pollinated weedy which, when covered with moist litter root, tle deaths (A$0.5m), 4.5% reduced per- lantanas was 36.7–48% in inflorescences send up vigorous shoots from the nodes. formance (A$2.0m), 7.3% loss of pasture averaging 32.6 flowers each (Auld 1969, These shoots may be confused with root (A$3.0m) and A$2.2m in control costs Mathur and Mohan Ram 1986, Swarbrick suckers. (Culvenor 1985). unpublished). Since there are normally After cutting back lantana regrows vig- Similar problems exist in other tropical two inflorescences per node during active orously from dormant buds at the bases and subtropical countries, e.g. in India growth, the total number of fruits pro- of the stems. It can also be propagated by (Singh et al. 1970) and (Water- duced by a lantana plant under good stem tip or hardwood stem cuttings house and Norris 1987). Annual losses growing conditions is several thousand placed in moist rooting media or by hard- due to lantana in the are per square metre per year. In the Philip- wood cuttings planted directly into moist estimated to be A$70 000 and in Fiji to be pines Cadigal (personal communication) soil. A$800 000 (Anon. 1993a). recorded an average of 24 fruits per inflo- rescence, 511 inflorescences per plant, Hybrids Natural . Lantana is a serious and 12 264 fruits per plant. The lantana complex is itself of invader of disturbed ecosystems con- Lantana seeds may either be trans- origin (e.g. Howard 1969), and hybridizes served for their natural and semi-natural ported away from the plant or fall beneath freely within itself (e.g. Howard 1970, attributes throughout its Australian and the parent plant. Most fruits are thought to Spies 1984) to produce fertile hybrid overseas distribution. It is among the be dispersed by birds, often locally but swarms morphologically similar to the most widespread and obvious of the ex- also to distances of up to 1 km or more. parent plants. Members of the lantana otic species in the Wet Tropics World Her- The seeds resist digestion during passage complex can also hybridize back to the itage Area of northern Queensland through the vector, and initial infestations original species from which the modern (Humphries and Stanton 1992), where it from bird-dispersed seeds are most com- complex arose. Sanders (1987) has invades disturbed rainforest. Whilst monly seen under fences, trees and other shown that a weedy tetraploid L. camara lantana persists along roadsides, creeks obvious perches. crosses with the diploid Floridan and other open situations it is shaded out L. depressa to produce a triploid resem- by the dense canopy as rainforest be- Physiology of seeds and germination bling the naturally occurring Bahaman comes re-established. Lamb (1981), on the Whilst investigating the effects of the L. ovatifolia, and that such hybrids form a other hand, has noted the persistence of lantana seed fly lantanae on the morphological continuum between the lantana for over 30 years after deliberate germination of lantana seeds from Natal, parental taxa. rainforest disturbance in south-eastern Graaff (1986) recorded a very low germi- Queensland. Lantana may play an impor- nation rate even in uninfested fruit. Of Importance tant role in the marginal fire ecology of 1200 seeds examined only 48% had fully rainforests, depending on local topogra- formed embryos. Breaking the dormancy Detrimental phy, wind and moisture content of the (possibly caused by the woody seed coat) Pastures. The main economic damage vegetation and admixture of other fuels by 180 days of dry storage increased caused by lantana is through decreased such as exotic grasses (Humphries and 87 Plant Protection Quarterly Vol.10(2) 1995 Stanton 1992). Under different conditions Table 1. Mean density of shrubs, saplings and trees per hectare in three lantana may either increase or decrease lantana free and lantana infested areas of woodland near Brisbane (Alcova the fuel load along the rainforest margin. 1987). Lantana is mainly an invader of open (in Australia especially eucalypt) wood- Mean density per hectare in each size class land (Lamb 1988, Humphries and Stanton (diameter 1m above ground) 1992). It commonly forms dense mono- <6 cm 6–15 cm 16–30 cm 31–60 cm >61 cm specific stands several metres tall which exclude native herbs, shrubs and tree and Lantana free 1248 436 173 85 14 climber seedlings, greatly reducing the Lantana infested 379 182 108 74 19 area’s conservation value, making it im- penetrable to people, and greatly increas- 1976/1977 (Master 1985) but has since giving it an advantage in competition ing the fire hazard under dry conditions. dropped to about A$0.5m (Wells 1984). with them once established (Swarbrick, Infested woodland near Brisbane had sig- Lantana control accounts for about 30% unpublished). Wadhawani and Bhardwaja nificantly fewer shrubs, saplings and trees of the establishment and 25–50% of the (1981) observed that lantana inhibits the in the smaller size classes in areas of dense harvesting costs of this crop (Wells germination of spores of the fern lantana than in adjacent and otherwise 1984a). Lantana is not a problem in es- Cyclosorus dentatus in India. Both similar lantana free areas (Alcova 1987) tablished exotic pine plantations which Achhireddy et al. (1985) and Singh et al. (Table 1). shade it out, but in 1966/1967 the Forestry (1989) have attempted to isolate the Dry lantana burns readily even when Commission of New South Wales spent allelopathic chemicals within lantana. green (Gujral and Vasudevan 1983), and about A$5500 attempting to establish bio- unless drought stressed the plant recov- logical control in indigenous forests within Host for other pests. In India lantana has ers quickly after fire from basal dormant the state (Baur 1967). Lantana is also a been shown to be a symptomless host for buds. Together with its smothering nature serious weed of forestry in India (Ghosh the sandalwood spike disease caused by this gives lantana a distinct advantage in 1980), ousting valuable timber species, Chlorogenus santali and a primary host for moist coastal and subcoastal woodlands. inhibiting natural regeneration of forest the dodder Cuscuta reflexa from which it Lamb (1988) showed that during the first trees, causing problems during the estab- spreads to more useful plant species ten years after invasion lantana altered lishment of plantations, creating a serious (Nayar and Srimathi 1968, Prasad 1966). the whole balance of eucalypt woodland. fire hazard and poisoning grazing stock. Waterhouse and Norris (1987) reported The amount of wood and twig litter fall that lantana provides cover for rats, wild decreased as the native trees lost vigour, Tropical Plantations. Lantana is a major pigs and other vermin in Fiji and provides whilst the amount of lantana litter fall weed of coconuts throughout the tropics, a favourable microclimate for in sharply increased. The amount of nutri- thriving in the moist disturbed soils and East Africa. ent in the litter decreased, suggesting that high light conditions, significantly reduc- more was taken up by the lantana with ing yields and interfering with manage- Beneficial less available to the native species. The ment and harvest (Thaman 1974, Ornamental. Both the weedy and non- concentration of nitrate nitrogen in the soil Waterhouse and Norris 1987). Annual weedy forms of lantana are grown exten- and litter increased to the advantage of production losses of 1–3% occur in coco- sively as garden ornamental shrubs in the the lantana and other exotic weeds. Lamb nut plantations in Vanuatu, Solomon Is- areas in which it is a weed and elsewhere. concluded that ‘the processes by which lands and Fiji, worth A$230 000, A$390 The non-prickly, non-fruiting triploid the community maintains itself ... are at 000 and A$500 000 respectively (Anon. forms make better ornamentals than the risk, and the longer the process continues 1993a). prickly fruiting forms, since they are the more unpredictable the result will be.’ shorter and denser in habit and their Regeneration of native plant communities Transport and industry. Lantana is one of flowers persist much longer due to fail- after clearing dense lantana may be slow the eight most troublesome weeds affect- ure of fertilization. where few viable native plant propagules ing railway land in New South Wales, be- are left in the soil. For the first year or cause of its size and the rapidity of its Essential oils and . The more such areas tend to be dominated by spread (Mahoney 1967). It is also a prob- mixed essential oils in lantana flowers ephemerals including cobbler’s pegs lem along disturbed power line and leaves can be fractionated to provide (Bidens pilosa), after which exotic and na- easements throughout south-eastern and oils useful for perfumery and possibly tive grasses begin to dominate and shrub coastal Queensland. beneficial drugs (Ahmad et al. 1962, and tree seedlings appear (Swarbrick un- Peyron et al. 1971), but do not seem to published). Allelopathy. Like most other plants, have been commercialized as yet. The Lantana is an aggressive invader of lantana can be shown to produce chemi- ability of lantana and its extracts to con- natural ecosystems in many other coun- cals which inhibit the germination and trol other organisms suggests that it may tries, for example, the Galapagos Islands growth of other plants in the laboratory. be used as a biocide. Dried and powdered (Washington 1985), many Pacific islands The allelopathic effect of lantana against lantana protects stored potatoes against (Thaman 1974) and South Africa (Stirton agronomic crops such as wheat and potato tuber in south-east 1978). soybeans and annual weeds (Mersie and (Anon. 1988), whilst lantana extract can Singh 1987) is probably of little impor- protect ripe tomato fruits against the Forestry. Lantana is a serious weed of tance, but Singh and Achhireddy (1987) combination of a fruit fly and an associ- commercial hoop pine plantations in have shown that lantana is also ated fruit rotting fungus (Sinha and coastal south-eastern Queensland, com- allelopathic to citrus, a crop in which it is Saxena 1987). On the other hand Ahmed peting with tree seedlings for light, inter- a major weed in Florida. Achhireddy and and Agnihotri (1977) have shown that fering with access throughout the life of Singh (1984) showed that lantana is lantana leaf extracts failed to inhibit a the plantation, and increasing both man- allelopathic to the perennial weed range of test fungi. agement and harvesting costs and fire Morrenia odorata in Floridan citrus or- hazard. The cost of lantana control in chards. Lantana may also be allelopathic Stockfeed. Lantana seeds are at times so these plantations exceeded A$0.9m in to many endemic Australian plants, abundant in India that they can be Plant Protection Quarterly Vol.10(2) 1995 88 Table 2. reported for the control of lantana. Method of application Foliar Soil Basal bark Cut Stump Stem injection Ground Aerial 2,4-D + 2,4,5-T 10,23,24,25 20 2,4-D Q,N,3,4,9,10,12, 2 Q,N,2 Q,N,13 16,18,19,23,25 2,4,5-T 11,23 Q 11 Q Amitrole 4 Bromacil 14 Buthidazole 2 Dicamba 20 Dicamba + MCPA 9,17,18,19 Dicamba + 2,4,5-T 11 Dichlorprop Q,N,2,3,6,9, 2 Q,2 Q 16,17,18,19 Ethidimuron 22 Fenoprop 22 20 Fluroxypyr Q,2,27 Fosamine 4,9,12,16,17,18,26 Glyphosate Q,N,SA,2,7,9,11,12, 2 8,11 17,18,19,22,26,29 Hexazinone 2 Imazapyr SA,7,11,19 SA,7,8,11 Metsulfuron methyl Q,2,8,9,19 18,21,27 Paraquat 24 20 Paraquat + diquat 20 Picloram SA,7,10,12 2 2 SA,7,13 Picloram + 2,4-D Q,N,9,12,17,18,19 20 13 Picloram + 2,4,5-T 10,11 11 Picloram + triclopyr Q,N,8,9,18,19 SA,7,8 Tebuthiuron Q,N SA,2,7,14 Triclopyr Q,11,26 Q Q Q,N,11,13 Q Registered for use on lantana in Queensland, Australia (Source: Infopest) N Registered for use on lantana in New South Wales, Australia (Source: Peskem) SA Registered for use on lantana in South Africa (Source: Erasmus 1991) 1 Abell 1973 2 Armstrong et al. 1987 3 Batholomew and Armstrong 1978 4 Campion 1978 5 Chang et al. 1982 6 Diatloff and Percy 1973 7 Erasmus 1991 8 Erasmus 1992 9 Fenton 1987 10 Ghosh 1980 11 Graaff 1986 12 Killilea 1983a 13 Killilea 1983b 14 Killilea 1983c 15 Love 1989 16 Master 1985 17 McMillan and Strachan 1984 18 McMillan and Strachan 1989 19 McMillan 1989a 20 Motooka et al. 1967b 21 Motooka et al. 1991 22 Phillips and Tucker 1976 23 Sinha 1976 24 Soni 1980 25 Tam 1947 26 Toth and Smith 1984 27 Vitelli and van Haaren 1988 28 Vitelli and Dorney 1991 29 Wells 1984a harvested and used for stockfeed (Lall this attempt by showing that even the (fluroxypyr, picloram, clopyralid and et al. 1983a and 1983b). triploid forms are capable of producing triclopyr) groups. Inhibitors of fertile pollen and occasional viable seeds acetolactate synthetase such as the Legislation and can hybridize with other forms of the sulfonylureas (metsulfuron methyl) and Lantana was a declared noxious weed plant. imidazolinones (imazapyr) and inhibitors throughout much of New South Wales for of EPSP synthetase in the shikimic acid many years, but was removed from the Response to herbicides pathway such as glyphosate also show noxious weeds lists in the 1980s since it Many chemical control trials have been good activity. Inhibitors of photosynthe- was uneconomic to control in most situa- conducted against the various forms of sis such as the triazine, uracil, urea and tions. Parsons and Cuthbertson (1992) re- lantana in Australia, South Africa, Zim- bipyridyl groups of herbicides have little ported that it is still a declared plant in babwe, India, Taiwan and the USA (Table effect on lantana, possibly due to its abil- certain shires and municipalities in New 2). The comparative efficacy of various ity to drop leaves and refoliate rapidly. South Wales and is a Classes B and C herbicides and factors affecting their effi- The order of decreasing effectiveness weed throughout the Northern Territory cacy are discussed below. on red flowered lantana in northern New outside town areas. South Wales was glyphosate, picloram/ Although all forms of lantana are de- Effect of herbicide type triclopyr, 2,4-D + picloram, dichlorprop, clared noxious weeds in Natal, attempts The most effective herbicide groups are fosamine, dicamba + MCPA, metsulfuron have recently been made to allow the sale the phenoxy acid (2,4-D, 2,4,5-T, methyl and 2,4-D (Fenton 1987). Trials of the largely sterile triploid forms. Spies dichlorprop and MCPA), benzoic acid over three years with ‘Helidon White’ indi- and du Plessis (1987) have reacted against based (dicamba), and pyridine based cated the following order of decreasing 89 Plant Protection Quarterly Vol.10(2) 1995 effectiveness at the rates used: fluroxypyr, between various surfactants used with mortality due to foliar applied glyphosate, dichlorprop and metsulfuron metsulfuron methyl in Hawaii (Motooka fluroxypyr, with plant mortality decreas- methyl (Hannan-Jones unpublished). et al. 1991). ing after a period with average minimum Fluroxypyr controlled ‘Common Pink’ temperature below 15°C and more than more effectively than metsulfuron methyl Effect of season and climate 30 mm of rain being required in the pre- (Vitelli 1991, Armstrong et al. 1987), whilst Control of lantana with herbicides shows ceding six weeks for a significant re- at the rates used fosamine was equally or a distinct seasonal response for sponse from foliar applied fluroxypyr more effective than the recommended glyphosate, fluroxypyr, dichlorprop and (Hannan-Jones unpublished). rates for 2,4-D amine or dichlorprop (Mas- metsulfuron methyl for the ‘Helidon White’ Low rainfall before cut stump treat- ter 1985). Toth (1984) demonstrated that form in south-eastern Queensland ment has reduced the effectiveness of glyphosate was more effective than (Hannan-Jones unpublished, Figure 4). many herbicides in Zimbabwe (Killilea fosamine, whilst Armstrong et al. (1987) This effect has been shown for these and 1983a). In Australia good results have showed glyphosate to be more effective other herbicides at other locations and for been obtained under wet summer condi- than dichlorprop as a low volume foliar other lantana forms (Killilea 1983b, Wells tions by Fenton (1987) and the best results spray with a sprinkler sprayer, particularly 1984b, Fenton 1987, Vitelli and Van for fosamine have coincided with the when applied in a wet autumn and under Haaren 1988). For growth regulating hor- main autumn flowering period (Campion shade. mones, such as 2,4-D, dichlorprop, 1978). Soil applied picloram granules, fluroxypyr and picloram, which are most tebuthiuron, hexazinone and buthidazole effective between December and April, Effect of plant form all failed to control lantana in south-east- this effect is closely connected to the The great variability within the lantana ern Queensland with or without prior burn- growth activity of the plant and possibly complex may well be responsible for its ing (Armstrong et al. 1987). In Hawaii, with increased translocation of the ap- variable response to similar herbicide however, hexazinone has been shown to plied herbicides within the plant. In the treatments (Graaff 1986). Some reports be as effective as tebuthiuron in some case of fosamine, the effect is associated indicate that 2,4-D is effective for lantana situations (Motooka et al. 1991). with the onset of leaf senescence and a control (Abell 1973), while others state Tebuthiuron was more effective in No- possible basipetal translocation of that it is ineffective (Tam 1947). vember than in May in Zimbabwe (possi- fosamine into the wood of the plant and Various forms of lantana throughout bly due to a lack of rainfall after applica- acropetal translocation into the next Queensland have shown quite different tion), whilst soil applied bromacil was in- year’s buds (Campion 1978). responses to 2,4-D amine as a high vol- effective (Killilea 1983c). In north-eastern Australia autumn is a ume overall spray, in that pink and white time of reducing rainfall, and both Fenton flowered forms both died within four Effect of application method and (1987) and Vitelli and Van Haaren (1988) months, but red forms recovered, normal addition of surfactants have noted that the poorer results in au- regrowth appearing within six weeks of The response of lantana to a given herbi- tumn correspond with water stress and treatment even though there were no sig- cide can be influenced by its method of reduced foliage cover, whilst Killilea nificant differences in either leaf application and by the addition of (1983b) has related this effect to the onset wettability or spray retention (Diatloff surfactants. of cooler weather. Trials in south-eastern and Haseler 1965). In Northern New High volume foliar application of Queensland have shown correlations be- South Wales McMillan (1989a) has shown fluroxypyr was a more effective method tween temperature and rainfall and plant that red-flowered lantanas are more than low volume gas gun, sprinkler sprayer or aerial application (Love 1989). 100 However, fosamine was more effective when applied by low volume gas gun than by high volume overall spraying (Toth and Smith 1984). Herbicide screening by the Queensland 80 Department of Lands on ‘Helidon White’ (Armstrong et al. 1987) indicated that 2,4-D was more effective than di- chlorprop when applied as a basal bark 60 treatment whereas the reverse was true when comparing their high volume foliar efficacy. In Hawaii, triclopyr was ineffec- tive as a foliar application, but was effec- 40 tive when applied as a basal application in diesel oil, which suggested that there was some impediment to triclopyr uptake % mortality (12 MAT) in foliar treatments (Motooka et al. 1991). 20 Aerial application of 2,4-D, glyphosate, dichlorprop and picloram has given unsatisfactory control of lantana, whereas ground application has been shown to be quite effective (Armstrong 0 et al. 1987). Nov-90 May-91 Nov-91 May-92 Nov-92 The addition of a surfactant to Date of application fluroxypyr gave better control of one undescribed form in northern Queens- Figure 4. Seasonal response of L. camara to selected foliar applied land (Love 1989), but there appeared to herbicides (Hannan-Jones, unpublished). be no significant differences in effect ● Fluroxypyr, ■ Glyphosate, ▲ Metsulfuron methyl, ◆ Dichlorprop. Plant Protection Quarterly Vol.10(2) 1995 90 difficult to control with herbicides than In south-east Queensland several trials sap sucking bug scrupulosa ‘Common Pink’. Only 2,4-D and are being conducted into the direct sow- have been released in the Solomon Is- fluroxypyr have exhibited variable effects ing of competitive fodder plants such as lands (Anon. 1993b) (Table 3). on lantanas of different flower forms in such as glycine (Neonotonia wightii) and The exploration for and primary host northern Queensland (Vitelli and Dorney leucaena (Leucaena leucocephala) into specificity testing of these insects has been 1991). lantana thickets (in the belief that grazing carried out by the Hawaiian Department In Mauritius the pink flowered form has stock attempting to graze will trample of Agriculture, the Commonwealth Sci- been shown to be easily killed by MCPA down the lantana) and into the replace- entific and Industrial Research Organiza- or 2,4-D, whereas even under the most fa- ment of lantana with native trees (M. tion and the Queensland Department of vourable conditions the lethal dosage for Roberts personal communication, J.C. Lands, Australia. The distribution of the orange flowered form is three times as Galletly personal communication). De- biocontrol agents to many African and great (Birch 1961). graded rainforest clearings in northern Pacific Island countries has been carried New South Wales choked with lantana out by the International Institute of Bio- Effect of plant size are also being regenerated by planting logical Control using species already im- In Queensland it has been observed that and tending selected rainforest pioneer, ported into other countries, and the South glyphosate gives decreasing control of secondary and tertiary species until a African Plant Protection Research Insti- lantana with increasing plant size de- canopy is formed and the lantana is re- tute has imported and released biocontrol pending on the method of application duced by a lack of light (R. Hynes per- agents in South Africa since 1961 (Cilliers (Wells 1984b), whilst fosamine is decreas- sonal communication). and Neser 1991). ingly effective as plant height exceeds The Queensland Forest Service has Despite this massive effort lantana re- 2 m, perhaps due to a ‘shading effect of successfully used a combined harvester mains one of the world’s worst weeds plant size on foliage coverage or ... larger and mulcher to clear lantana and other (Holm et al. 1977) and thus its biological plants [having] a physiological character- woody regrowth from between rows of control can be considered to have been istic that made them less susceptible to hoop pine (Araucaria cunninghamii) plan- unsuccessful, even though levels of con- fosamine’ (Master 1985). Similarly Killilea tations to allow access to the trees for trol ranging from minor to significant have (1983b) noted that smaller plants were trimming and felling, but the machine is been achieved in some countries. The more easily killed than larger bushes by unsuitable for rough or hilly terrain (P. main reasons for this overall lack of suc- overall spraying with a number of herbi- Lees personal communication). cess can be attributed to the extreme vari- cides and that plants with thinner stems In India the establishment of fodder ability of the plant coupled with its ability were more susceptible to cut stump treat- trees such as Grewia optiva, Bauhinia to colonize diverse habitats. Introduced ment. The volume of applied herbicide variegata and Populus spp. after clearing insects may show preferences for one or required to control lantana has been lantana has shown some success in pre- more of the many forms of lantana or may shown to be proportional to the volume venting the return of lantana. The fast fail to thrive in some of the environments of the plant rather than the leaf surface spreading vine Pueraria thunbergiana is in which the weed can survive. area (McMillan 1992), which may explain also a promising replacement for lantana The insect species used as biological some of the size effects described by ear- (Sharma 1988). In Zambia, Anderson control agents have been collected from lier workers. (1963) investigated the clearing of lantana several species of Lantana. These include from amongst thick natural bush using a L. tiliaefolia and L. glutinosa in Brazil Response to fire, physical control combination of trimming and specially (Winder and Harley 1983) (both now con- and competition designed one- or three-man operated le- sidered to be subspecies of L. urticifolia, Although the use of fire to control of vers to grub out individual lantana Palmer and Pullen in press), and lantana in pasture is widely recom- bushes. All but the longest lateral roots L. camara, L. urticifolia, L. urticoides and mended (e.g. Goodchild 1951, Saint- can be completely removed, but soil dis- L. hirsuta in Mexico, USA and Central Smith 1964, Bartholomew and Armstrong turbance causes a flush of lantana and America (one or more of which belong to 1978), very little experimental work has other weed seedlings. the camara section of the genus, Palmer been published on its use and effects. Fire and Pullen in press). Most agents have is generally insufficient by itself for Response to natural enemies been collected from Mexico, with the only lantana control, and additional treat- Surveys of the phytophagous insect fauna successful agent from Brazil being ments including sowing competitive pas- of Lantana camara and closely related spe- U. girardi (Table 3). ture species, controlled grazing and fol- cies have yielded 544 species in North and The successful establishment rate of low-up burns and chemical treatments Central America (Palmer and Pullen in control agents has ranged from zero with are usually required. Little information is press) and 345 species in Brazil (Winder Alagoasa parana, Autoplusia illustrata and available on the seasonality of burning, and Harley 1983). Most are probably Teleonemia elata to over 95% with the effect of meteorological conditions at polyphagous and thus unsuitable as bio- U. girardi, Epinotia lantana and the time of the burn or the occurrence of logical control agents. T. scrupulosa, whilst their effectiveness rain after burning, the quality and type of Lantana was the first weed against has ranged from established but having fuel, whether a ground or crown fire is which biological control was attempted. no discernible control through minor and preferable, or time of reintroduction of Koebele in 1902 collected 23 insect species partial to significant control (Julien 1992) stock after the fire. in Mexico and shipped them to Hawaii, (Table 3). Several authors have reported the ef- where eight were eventually established Studies of the effect of introduced in- fects of mechanical clearance of lantana in (Perkins and Swezey 1924). Julian (1992) sects on lantana have been limited to the pastures (e.g. Goodchild 1951, Saint- lists 32 insect species that have been four species most widely established, Smith 1964, Bartholomew and Armstrong deliberately released in 25 countries. Since , Teleonemia scrupulosa, 1978). All have shown that cultivation then the leaf-feeding chrysomelid Uroplata girardi and Octotoma scabripennis. gives very effective control of lantana Charidotis pygmaea and oecophorid moth The agromyzid fly Ophiomyia lantanae but that effective pasture establishment Ectaga garcia have been released in was one of the original insects found on and management are essential and that Queensland (Willson 1993) and the leaf- lantana by Koebele and is now estab- some follow-up control is generally re- mining chrysomelid Uroplata lished in nine of the 11 countries in which quired. girardi and Octotoma scabripennis and leaf it has been released. It oviposits in green 91 Plant Protection Quarterly Vol.10(2) 1995 Table 3. Insect species released as biological control agents of Lantana camara, their establishment and effectiveness. Species Country of origin of agent Lantana hostA No. of countries Effectiveness ratingB released established Root Feeders Coleoptera Parevander xanthomelas Mexico C 1 0 – Stem Feeders Coleoptera championi Mexico C,H,U 1 0 – Plagiohammus spinipennis Mexico H 5* 2* 1–3 Hepialus sp. Mexico – 1 0 – Diptera Eutreta xanthochaeta Mexico C,U 3* 1 1 Leaf Feeders Leptobyrsa decora Peru and Colombia – 9* 2* 1–2 Teleonemia elata Brazil T,G 5* 0 – Teleonemia harleyi Trinidad – 1* 1* 1 Teleonemia prolixa Brazil T,G 1* 0 – Teleonemia scrupulosa Mexico C,H,U,Uc 21* 20* 1–3 Coleoptera Alagoasa parana Brazil T,G 2* 0 – Charidotis pygmaea Brazil T 1* 0 – Octotoma championi Costa Rica C,H,U 3* 1* 1 Octotoma plicatula Mexico – 1 0 1–4 Octotoma scabripennis Mexico C,U 9* 6* 2 Uroplata fulvopustulata Mexico, Costa Rica, Panama C,U 3* 1* – Uroplata girardi Brazil T,G 21* 19* 1–4 Uroplata lantanae Brazil T 2* 0 – Lepidoptera Autoplusia illustrata Colombia 2* 0 – Cremastobombycia lantanella Mexico C,H,U,Uc 1 1 2 Panama, Trinidad U 10* 1 1 Ectaga garcia Brazil T 18 0 – strigata Kenya, Zimbabwe – 6* 6* 1–2 Neogalea esula C,U,Uc 4* 2* 1–2 Pseudopyrausta acutangulalis Mexico C,U,Uc 3 0 – haemorrhoidalis Florida, USA, Cuba C 12* 7* 1–3 Diptera Calcomyza lantanae Trinidad C 2* 2* 2–3 Flower Feeders Lepidoptera Epinotia lantana Mexico C,H 4* 4* 1–3 Lantanophaga pusillidactyla Mexico C,H 4 3* 1–3 Thecla bazochii Mexico C,H 3* 2 1 Thecla sp. Mexico C 2 1 1 Seed/Fruit Feeders Coleoptera Apion sp. A Mexico T 1 0 – Apion sp. B Mexico – 1 0 – Diptera Ophiomyia lantanae Mexico – 11* 9* – A Insects found on the following species: C=L. camara, H=L. hirsuta, U=L. urticifolia, Uc=L. urticoides, T=L. tiliaefolia, G=L. glutinosa. B Effectiveness rating: 1 = established, 2 = minor control, 3 = partial control, 4 = significant control. * Released and/or established in Australia. fruits, where larvae feed on the fleshy less attractive to birds, slowing down the significantly reduced by the fly, contrib- pulp and may enter the seed and feed on rate of spread of lantana. Indian studies uting to lantana control (Graaff 1987). the embryo (Thakur et al. 1992). Larvae by Thakur et al. (1992) showed that the The tingid bug T. scrupulosa is estab- pupate either in the fruit or peduncle insect has little effect on germination, lished in 20 countries. It oviposits in (Cilliers and Neser 1991), and both which in both infested and non infested stems, petioles and peduncles, most often Wilson (1960) and Thakur et al. (1992) seed was less that 15%. In South Africa where adults have flowers on which to have shown that they make the berries seed germination at some sites was feed (Currie and Fyfe 1939). The nymphs Plant Protection Quarterly Vol.10(2) 1995 92 feed in groups (generally on the under- attacked the roots. The root boring are similarly damaging in tropical areas. sides of leaves) and the adults on leaves, cerambycid beetle Parevander xanthomelas Laboratory and greenhouse studies have flowers and occasionally young stems. was released in Hawaii in small numbers commenced to determine their biology Oviposition and feeding cause chlorosis, in 1902 but failed to establish. This beetle and immediate host range (Tomley and distortion and leaf abscission, heavy has now been approved for introduction Evans 1992). attack resulting in complete defoliation into Australia for host specificity testing and systemic dieback of the plant (Harley in quarantine. Acknowledgments and Kassulke 1971). The bug prefers low The stem boring cerambycid beetle The authors wish to thank G.G. White for rainfall areas, and populations peak dur- Aerenicopsis championi was described by helpful comments on the manuscript. ing autumn and early winter before Mann (1954) as an insect of great poten- drought and frost defoliate plants, caus- tial, but failed to establish in Hawaii de- References ing insect numbers to fall rapidly. Harley spite large numbers being released Abell, T.M.B. (1973). The value of herbi- et al. (1979) suggest that population levels (Fullaway 1958). It has been host tested cides to forestry in Zambia: a survey of may be influenced by changes in the in Australia and is expected to be released work to date by the research division. physiology of the plant. Randomized ex- in Queensland in 1994. Eggs are laid in Forest Newsletter (Zambia Forest De- clusion trials by Harley et al. (1979) leaf petioles and the larvae on hatching partment) 120, 43-5. showed that a red flowered form was the feed down through the petioles into the Achhireddy, N.R. and Singh, M. (1984). preferred host, with ‘Common Pink’ be- tips and then down the stems for dis- Allelopathic effects of lantana (Lantana ing the least preferred, whilst in South tances of up to 1.2 m. As the larvae feed, camara) on milkweed vine (Morrenia Africa Cilliers (1987) has noted that the stems are hollowed out and killed. odorata). Weed Science 32, 757-61. T. scrupulosa has also shown a preference Another stem boring cerambycid Achhireddy, N.R., Singh, M., Achhireddy, for certain forms of lantana. Heavy attack Plagiohammus spinipennis has been re- L.L., Nigg, H.N. and Nagy, S. (1985). by the bug in Queensland helps to reduce leased in five countries, but has only be- Isolation and partial characterization of the aggressiveness of some forms of come established in Hawaii where it is ex- phytotoxic compounds from lantana lantana, and although it is common erting some control in high rainfall areas. (Lantana camara L.). Journal of Chemical throughout lantana areas of South Africa The eggs are inserted into the stems usu- Ecology 11, 979-88. it seldom reaches highly damaging popu- ally within 600 mm of the plant base. The Ahmad, M.N., Bhatty, M.K. and lation levels. larvae girdle the stems and tunnel into the Karimullah (1962). Some essential oil U. girardi and O. scabripennis are leaf- xylem tissue. Multiple infestations of sources from West Pakistan. Pakistan mining hispine beetles. O. scabripennis is stems result in them being killed or weak- Journal of Science 14, 12-5. established in six countries and U. girardi ened so severely that they are readily bro- Ahmed, S.R. and Agnihotri, J.P. (1977). in 19. The eggs are laid singly into the ken, reducing tall plants to mutilated Antifungal activity of some plant ex- upper surface of leaves and covered with stumps (Harley 1969b). Although estab- tracts. Indian Journal of Mycology and an exudate from the insect’s collaterial lished in low numbers at one site in Aus- Plant Pathology 7, 180-1. glands (Harley 1969a). The larvae tunnel tralia (Taylor 1989), P. spinipennis has Alcova, P.A. (1987). The effects of the into the leaf, feeding on the mesophyll and failed due to the plant forming callus tis- presence of Lantana camara on local bird leaving both upper and lower epidermis sue at the oviposition site, destroying the populations in Brisbane Forest Park. undamaged whilst forming feeding gal- eggs and neonate larvae before they enter Project Report No T(AS)99. (Queens- leries radiating from a central chamber the xylem tissue (Willson 1974). It appears land University of Technology, Bris- before pupating in the leaf. The resultant that the success of P. spinipennis in Hawaii bane). mines of O. scabripennis are much less tor- was due to the condition of the stems that d’Almeida, C. and Roland-Heydacker, F. tuous that those of U. girardi. After emer- were very soft and spongy near to the (1985). Etude comparative du pollen de gence the adults feed on the upper leaf ground (K.L.S. Harley personal commu- Lantana camara L. () surface. The effect of these two beetles nication). A similar condition prevailed in originaire du Benin et de Martinique. on lantana has been studied by Cilliers Mexico where Koebele described the Pollen et Spores 27, 321-34. (1987), who showed that in association lantana stems as usually very soft and Anderson, P.E. (1963). Lantana eradica- with T. scrupulosa they destroy 25% of the knotty near the ground as a result of a tion by manual means. Bulletin 2195, annual leaf production of lantana in Na- fungal disease (Perkins and Swezey 1924). 95-6. tal. A stem sap sucking membracid bug, Anon. (1988). Lantana controls insects. O. scabripennis prefers cooler inland ar- , is currently under- Tropical Weed News No 3. Long eas whereas U. girardi does well in hot going host specificity tests in Queensland Ashton Research Station, Bristol, UK. wet areas. In Queensland these insects and is expected to be released within 12 Anon. (1993a). Economic impact of pests can cause defoliation of lantana over the months. This insect causes considerable and weeds. SPC Agricultural News, summer–autumn months, markedly re- dieback of lantana stems in Mexico Newsletter of the South Pacific Commis- ducing its vigour, competitiveness and (Willson and Palmer 1992). sion Agriculture Programme 2(1), 9. flowering. This defoliation allows grasses Other insect species under investiga- Anon. (1993b). Successful biocontrol of to grow through the lantana plants, which tion by Queensland Department of Lands lantana. SPC Agricultural News, News- are further damaged if grazed by cattle. A are the leaf-mining gracillarid moth letter of the South Pacific Commission Ag- cold climate strain of U. girardi has been Cremastobombycia lantanella and the leaf- riculture Programme 2(2), 1. established in inland areas of South Af- sucking mirid bug Adfalconia intermedia. Armstrong, T.R., Harvey, G.J. and Vitelli, rica (Cilliers and Neser 1991), and after its Four fungal pathogens have been J.S. (1987). Lantana chemical control, release in the Solomon Islands in 1993 found in Brazil. These are the rusts herbicide screening. In ‘1986–87 Annual U. girardi successfully controlled the ‘Ha- Prospodium tuberculatum and Puccinia Report of the Biological Branch, waiian Pink’ form on an island in the lantanae, a hyphomycete Mycovellosiella Queensland Department of Lands’, p. 6. Russell Islands group (Anon. 1993b). lantanae and Ceratobasidium sp., a web (Queensland Department of Lands, While T. scrupulosa, U. girardi and O. blight. Field assessments indicate that Brisbane). scabripennis have been effective in attack- P. tuberculatum and M. lantanae are both Auld, B.A. (1969). The Ecology of Major ing leaves and tips of lantana, none of the damaging and host specific in cool cli- Woody Weeds of the Far North Coast agents released to date have successfully mates. P. lantanae and Ceratobasidium sp. of New South Wales. M.Sc. thesis, 93 Plant Protection Quarterly Vol.10(2) 1995 University of Sydney. Varietal differences in lantana. Pro- for reducing germination in Lantana Bailey, F.M. (1897). On some of the intro- ceedings of the First Australian Weeds camara forms found in Natal. South Af- duced plants of Queensland. Proceed- Conference, Toowoomba, pp. 1-11. rican Journal of Botany 53, 104-7. ings of the Linnaean Society of New Diatloff, G. and Percy, L.J. (1973). A new Gujral, G.S. and Vasudevan, P. (1983). South Wales 4, 26-36. formulation for lantana control. Austral- Lantana camara L., a problem weed. Bailey, J. (1841). Plants in flower in the ian Weeds Research Newsletter 18, 18. Journal of Scientific and Industrial month of June. (A list of plants flower- Driscoll, P.V. and Quinlan, G. (1985). Uti- Research 42, 281-6. ing in the Botanical Gardens of South lization of Lantana camara by birds and Harley, K.L.S. (1969a). The suitability of Australia in June 1841). South Austral- small mammals in Brisbane Forest Octotoma scabripennis Guer. and ian Magazine 1, 30. Park. Proceedings of a Workshop on Uroplata girardi Pic (Col., Bartholomew, B.L. and Armstrong, T.R. Resource and Recreation Management Chrysomelidae) for the control of (1978). A new look at lantana control. in Brisbane Forest Park. (Queensland lantana (Verbenaceae) in Australia. Queensland Agricultural Journal 104, Institute of Technology, Brisbane). Bulletin of Entomological Research 58, 339-44. Dronamraju, K.R. (1958). The visits of in- 835-43. Baur, G.N. (1967). Economics of weeds in sects to different coloured flowers of Harley, K.L.S. (1969b). Assessment of the forestry. Proceedings of the Weed Soci- Lantana camara. Current Science 11, suitability of Plagiohammus spinipennis ety of New South Wales 1, Paper 6, 452-3. (Thoms.)(Col., Cerambycidae) as an 3 pp. Erasmus, D.J. (1991). Control of Lantana. agent for control of weeds of the genus Birch, E.B. (1961). 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