Invasion, establishment and spread of Lantana (Lantana camara) - key threatening process listing

NSW Scientific Committee - final determination

The Scientific Committee has found that:

1. Lantana (Lantana camara L. sens. lat., family Verbenaceae) is a profusely branching, scrambling aromatic shrub 2-4 (up to 8) m tall and wide, with square-sectioned, often prickly cane-like stems. The leaves are opposite, ovate, often toothed, 2-6 (up to10) cm long. Flowers are 20-40 together in clustered compact heads 2-3 cm diam.; flower colour is variable, from cream and yellow to orange, red, pink, and mauve; flowers usually open yellow and change colour as a signal to pollinators. Flowering occurs through most months of the year, in NSW especially from September to April. The fruit is a single-seeded 'berry' (drupe) 4-8 mm diam., fleshy and purplish-black when ripe. Some variants are non-fruiting. Full descriptions are available in Swarbrick et al. (1995), Parsons and Cuthbertson (2001), and - for variants within the species in Australia - Smith and Smith (1982) and van Oosterhout (2004).

2. L. camara is an 'aggregate species', or 'species complex'. There are several natural variants of L. camara across its presumed native range in the tropical Americas, and in addition some hundreds of horticultural colour and habit varieties have been developed around the world, with over 650 varietal names coined (Howard 1969; Smith and Smith 1982). Some variants have arisen as a result of hybridisation with species in other related complexes such as L. urticifolia Miller (Day et al. 2003). Some variants may have hybridised with each other to form new feral wild-types. There are 29 morphologically defined variants of L. camara generally accepted to be naturalised in Australia (Smith and Smith 1982; Day et al. 2003), and new forms or biotypes may be evolving. Research to correlate morphological variants with genetic and ploidy variation is needed for a full understanding of the taxonomic variation. Taxonomy also needs to be correlated with life cycle traits, ecology and response to natural processes and management actions. The Australian variants differ in morphology and distribution, and in as yet imperfectly understood aspects of habitat preference, rates of fruit set, biocontrol susceptibility, response to different herbicides, and weediness.

3. Synonyms that have been applied to variants occurring in Australia include Camara vulgaris Benth., Lantana aculeata L., Lantana antidotalis Schumach. & Thonn., Lantana camara var. aculeata (L.) Moldenke, Lantana camara L. var. camara, Lantana camara var. crocea (Jacq.) L.H. Bailey, Lantana camara var. flava (Medik.) Moldenke, Lantana camara var. hybrida (Neub.) Moldenke, Lantana camara var. mista (L.) L.H. Bailey, Lantana camara var. mutabilis (Hook.) L.H. Bailey, Lantana camara var. nivea (Vent.) L.H. Bailey, Lantana camara var. sanguinea (Medik.) L.H. Bailey, Lantana camara var. splendens (Medik.) Moldenke, Lantana crocea Jacq., Lantana hirsuta M. Martens & Galeotti, Lantana mista L., Lantana nivea Vent., Lantana nivea var. mutabilis Hook., Lantana sanguinea Medik., Lantana scabrida Sol., Lantana scandens Moldenke, Lantana splendens Medik., and Lantana tiliaefolia Cham.

4. The basic chromosome number of L. camara sens. lat. is 11. Variants naturalised in Australia are mainly tetraploids, with some triploids, one diploid and one pentaploid (RJF Henderson, cited as pers. comm. 1971 by Everist 1974). Ploidy levels do not in general seem to be a barrier to interfertility (Day et al. 2003). Ploidy level does not appear to correlate in any straightforward manner with ecological characters (Day et al. 2003) or with morphology or toxicity (Henderson pers. comm., in Everist 1974). Nevertheless, the genetic and chromosomal levels of diversity within the species, its propensity for somatic mutation (Smith and Smith 1982, Day et al. 2003), and the interfertility of many of the variants, give it a high level of potential adaptability as a pest species. Some variants are putatively fruit sterile (and are sometimes sold as environmentally benign), but most or all may produce small amounts of viable fruit, and all are probably capable of contributing viable pollen to other variants, both factors increasing the likelihood of further feral genotypes (Spies and du Plessis 1987; Neal 1999, both cited in Johnson, (unpublished).

5. L. camara is native to Central and tropical South America. The earliest record for Australia is from 1841 in the old Botanic Gardens in Adelaide, and cultivated plants are known to have been grown in NSW by John Macarthur at Camden Park in 1843. There have been multiple introductions since, mainly in NSW and Queensland. It was reported by 1879 as a "most troublesome weed" in Queensland and "equally abundant all round Port Jackson" (Bailey 1879).

6. L. camara has spread along the east coast of Australia, from southern NSW north to Cape York, and from sea-level up to 600 m altitude, or less commonly to 1000 m. It has invaded at least 4 million hectares, mainly in NSW and Queensland (CRC Weed Management website), apparently based partly on Culvenor (1985)). In NSW most infestations are north of the Clyde River, with occasional mostly small occurrences from there south to the Victorian border; one of these however, at Mt Dromedary (Gulaga National Park and surrounds), is large. L. camara is also naturalised in the Northern Territory, South Australia and Western Australia, on Lord Howe and Norfolk Islands, and (apparently marginally) in Victoria. L. camara is not known to yet be naturalised in Tasmania, but its importation to that State is prohibited and a Statutory Weed Management Plan has been prepared. L. camara is naturalised in more than 60 other countries including New Zealand (Day et al. 2003).

7. On mainland Australia, L. camara has potential for much denser infestation of the coast and ranges, and to expand its range west and south of the Great Divide in NSW and other eastern States (Day et al. 2003), at least along creek lines. Where rainfall is the main determinant of soil moisture, the 750 mm isohyet has been suggested as an approximate lower limit for tropical Australia, and 650 mm in south-east Queensland (Swarbrick et al. 1995). The ecological constraints on spread southwards and to higher altitudes are uncertain, with some authors (Conn 1999; Parsons and Cuthbertson 2001; Day et al. 2003) suggesting that frost frequency and severity and/or generally lower temperatures are limiting factors. In contrast, Carr (1993) suggests the rarity of naturalisation in Victoria may be due to interacting climatic variables impinging specifically on germinant and seedling stages (with summer temperatures needed to trigger germination but summer rainfall patterns rarely providing sufficient moisture for successful germination and seedling establishment).

8. L. camara typically occurs where there is a moderate to high summer rainfall and well-drained sloping sites. Most variants have a preference for fertile organic soils, but some or all can survive on siliceous sands and sandstone-derived soils where these are of moderate depth and other conditions, especially year-round moisture, are suitable. L. camara does not tolerate waterlogging, salinity, prolonged drought, dense shading by overstorey species, frequent or severe frosts, or winter temperatures with prolonged periods < 5o C.

9. L. camara readily invades disturbed sites and communities. Various types of sclerophyll woodlands, sclerophyll forests, rainforests and dry rainforests are all susceptible to Lantana establishment (Driscoll and Quinlan 1985; Lamb 1988; Fensham et al. 1994; Gentle and Duggin 1997a), although in communities with a naturally dense canopy, Lantana colonisation may be heavily dependent on, and limited to, disturbance zones, edges, and canopy breaks. There is a strong correlation between Lantana establishment and disturbance (Stock and Wild 2002; Stock 2004), with critical factors being disturbance-mediated increases in light and available soil nutrients (Gentle and Duggin 1998) and, in rainforest, the competitive advantage of seedlings relative to many native species (Stock 2004). There is however considerable debate over the persistence and invasive capability of Lantana in relation to relatively undisturbed dense-canopy vegetation. Swarbrick et al. (1995, citing Swarbrick unpublished, and Humphries and Stanton 1992), argue that Lantana is "unable to invade tropical rain forest … [and] … dry vine scrubs" and "only persists along edges and where the canopy is broken", a conclusion broadly supported by Stock and Wild (2002) and - for subtropical rainforest - Stock (2004). In contrast, Fensham et al. (1994) postulate a probable role in dry rainforest for feral pigs causing initial canopy openings that allow Lantana establishment; pigs then avoid dense Lantana thickets, causing further openings in intact forest; the resulting Lantana thickets then, in this case, promote intense fire. In open forests and woodlands, given suitable soils and moisture, L. camara often becomes a dominant understorey species. In warmer, moister areas Lantana often becomes dominant in regenerating pastures. Lantana typically forms dense thickets, suppressing less competitive native vegetation and seedlings through shading (Swarbrick et al. 1995, ARMCANZ ANZECC&FM 2001), surface-soil nutrient sequestration (Lamb 1988 cited in Swarbrick et al. 1995; Gentle and Duggin 1998; CRC Weed Management 2003), smothering ("strangling" - ARMCANZ ANZECC&FM 2001) and perhaps through allelopathy (Gentle and Duggin 1997b; Day et al. 2003). Lantana provides habitat and food (fruits) for many species, including declared vermin and other pests, but also for native species, especially birds. It may also suppress other weeds, and management of the species must take account of these ecological roles. It can itself be suppressed eventually by shading following re-establishment of a sufficiently dense canopy (Fensham et al. 1994; Gentle and Duggin 1997a; Stock 2004). Lantana's effects on fire regime may vary considerably with site, and increased fire frequency and intensity will not always result from invasion by Lantana (Fensham et al. 1994; Stock 2004).

10. Lantana is pollinated by insects, mainly butterflies, thrips, and possibly some bees. There are conflicting reports about Lantana's ability to self-pollinate (Day et al. 2003). Neal (1999) reports 65% pollen fertility in one common weedy variety in Australia, but pollen and seed fertility probably differ significantly between variants. Fruit-set rates in weedy forms range from 37% to 85% (Swarbrick et al. 1995; Hilje 1985).

11. Seed dispersal is primarily by fruit-eating birds and to a lesser degree by foxes and other vertebrate foragers (Day et al. 2003). Seed longevity in the soil is not well documented, but 50% seed viability after 6 months dry shelf-storage has been recorded (ARMCANZ ANZECC&FM 2001), and seeds "are thought to remain viable for several years under natural conditions" (CRC Weed Management 2003). Work in progress (G. Vivian-Smith pers. comm.) suggests an in-soil longevity of at least three and up to five years, with significant variation by biotype. Germination rates are reported as being increased by removal of fruit pulp, as occurs with passage through bird gut (Day et al. 2003, CRC Weed Management 2003), and by warm temperatures, light, and high soil moisture. Vivian-Smith et al. (2006) however report that responses to fruit and seed damage are biotype-dependent and in some cases seedling emergence improves with seed damage. Germination rates even under favourable conditions are sometimes reported as fairly low (<45% or less), but as fruits may set at rates of up to several thousand /m2 there is a considerable soil seedbank. Gentle and Duggin (1997a) report that in-situ "burning, biomass removal, and soil scarification, either singly or in any combination, significantly increased germination", and that most combinations of these disturbances, all associated with cattle grazing and fire, favoured Lantana seedling growth rates.

12. L. camara is tolerant of occasional fire, occasional frost, and mechanical damage to the aerial stems, being capable of resprouting vigorously from the stem-base and of 'layering' (i.e. vegetative propagation by development of roots from stems in contact with soil). Individual plants may be very long-lived. The ability of Lantana to sucker from broken roots has been debated, with Swarbrick et al. (1995) asserting that it "does not sucker from damaged or broken roots, but will regrow vigorously from the base of the stem and more slowly from rooted horizontal stems in contact with moist soil", suppported by Stock (2004) and van Oosterhout (2004). In contrast, Parsons and Cuthbertson (2001) assert that the shallow lateral roots "sucker if damaged or broken", and "new canes are produced on the crown and from the lateral roots in early spring" S. Johnson (NSW Dept of Primary Industries unpublished, cites B. Johnson (pers. comm.)) as estimating that 30% of new shoots found after clearing of a site in northern NSW were of root-suckering origin.

13. Most variants of L. camara in Australia are toxic to domestic livestock (sheep, cattle) to some degree, with only three thought to be consistently non-toxic (Everist 1974, Parsons and Cuthbertson 2001). Toxicity seems likely to be related to genetic factors, not environmental ones (Everist 1974, citing Seawright 1965 and unpublished work by L.S. Smith). Toxins may occur in leaves, flowers, fruits and sap, and include triterpene acids (lantadenes A and B) and their reduced forms. Some toxic reactions have been recorded in humans, especially children. Palatability and toxicity to native herbivores do not appear to have been documented.

14. L. camara may change soil microhabitat through shading, self-mulching, and altered water and nutrient balances. Lamb (1988, cited in Swarbrick et al. 1995) identified an increase in soil nitrate in eucalypt woodland following Lantana invasion, to the benefit of the Lantana and other weeds, and to the detriment of some native species, and a decline in other nutrients. Gentle and Duggin (1998) point to Lantana's ability to aggressively compete for and sequester surface-soil nutrients, such as are made available by disturbance episodes, and verified experimentally Lantana's ability to out-compete and suppress an analogous native coloniser of mesic forests (Choricarpia leptopetala, Myrtaceae).

15. Lantana dominance may adversely affect the richness of some soil faunal assemblages. Cummings (2004) reports a reduced presence in Lantana-dominated vegetation of several functional groups of ant species, compared to adjacent 'natural' (rainforest and sclerophyll) communities, and concludes this probably relates both to initial disturbance events and to the structure of Lantana-dominated vegetation that followed. Overseas research (Fernandes et al. 2001) points to a progressive loss of diversity among mycorrhizal arbuscular fungi in rainforest systems of Madagascar, correlated with a loss of vascular plant diversity and non-autochthonous secondary successions including Lantana camara-dominated associations. Lantana is not established as a causal mechanism but the ability of Lantana to arrest vegetation succession for decades (e.g. Lamb 1994, apropos subtropical rainforest), and to inhibit growth of at least some microorganisms (Parsons and Cuthbertson 2001), suggests it may also inhibit mycorrhizal recolonisation.

16. The generally suppressive effect of Lantana on a wide range of native species is attested by several studies (Gentle and Duggin 1998, Day et al. 2003) and a multitude of field observations. Swarbrick et al. (1995), citing observations by Driscoll and Quinlan (1985) that "eucalypt seedlings generally fail to establish under lantana", infer inhibition of germination through lack of light. Webb et al. (1972) reports Lantana blocking the succession of rainforest species for some 15 years in an artificially created (1957) gap in intact rainforest, and Stock (2004) affirms that the site was still Lantana-dominated in 2004. Swarbrick et al. accept that Lantana may form a stable "community" arresting rainforest regeneration in some circumstances, but also assert that "many rainforest species germinate and develop under Lantana, eventually growing through it and shading it out", and Fensham et al. (1994) note the persistence of at least some regenerant dry rainforest tree species beneath Lantana thickets. Stocker and Mott (1981) note Lantana's suppression of grass invasion of rainforest margins, and its protective role against low-intensity early-season fires. It seems likely that rate of rainforest species regeneration (if any) through Lantana thickets will be determined by a number of factors including soil, rainfall, nature and repetition of disturbance, and local floristics (Stock 2004). L. camara is also thought to be allelopathic, i.e. able to inhibit or suppress by chemical means the germination and/or growth of at least some competing plant species. Overseas glasshouse studies (Swarbrick et al. 1995; Day et al. 2003) and a substantive Australian field trial (Gentle and Duggin 1997b), provide support for this.

17. Fensham et al. (1994) document declines in plant species richness with increasing levels of Lantana infestation of dry rainforest, and accumulation of heavy fuel loads along boundaries between savanna woodland and dry rainforest, leading to significant canopy tree loss and edge erosion in the latter, with Lantana dominating the area of lost dry rainforest and rendering it prone to further fires. Swarbrick et al. (1995) present summary data from Alcova (1987) showing a large (at least 70%) decline in inferred recruitment (number of native tree and shrub saplings present) in Lantana-infested areas of eucalypt woodland compared to Lantana-free areas.

18. In Queensland, L. camara has been identified as a potential threat to more than 60 plant and animal species of conservation significance (ARMCANZ ANZECC&FM 2001).

19. In New South Wales, Lantana has been identified (P. Downey pers. comm.) as a threat to the following native plant species listed on Schedule 1 and Schedule 2 of the Threatened Species Conservation Act:

Lantana may prove to be a threat to additional plant species or populations listed under the Threatened Species Conservation Act.

20. Lantana has been identified as a threat to at least two animal species listed as Endangered on Schedule 1 of the Threatened Species Conservation Act:

• Dasyornis brachypterus (Eastern Bristlebird)
• Ocybadistes knightorum (Black Grass-Dart Butterfly)

Lantana may prove to be a threat to additional animal species or populations listed under the Schedules of the Threatened Species Conservation Act. It adversely affects the ability of Koalas to move between trees (J. Shields pers. comm.)

21.Lantana has also been identified as a threat to the following Endangered Ecological Communities listed on Schedule 1 Part 3 of the Threatened Species Conservation Act (P. Downey pers. comm., D. Keith pers. comm.):

• Eastern Suburbs Banksia Scrub in the Sydney Basin Bioregion
• Illawarra Subtropical Rainforest in the Sydney Basin Bioregion
• Lowland Rainforest on Floodplain in the NSW North Coast Bioregion
• Littoral rainforest in the NSW North Coast, Sydney Basin and South East Corner Bioregions
• Lower Hunter spotted gum - Ironbark forest in the Sydney Basin Bioregion
• Pittwater Spotted Gum Forest
• River-flat eucalypt forest on coastal floodplains of the NSW North Coast, Sydney Basin, and South East Corner Bioregions
• Swamp Oak floodplain forest of the NSW North Coast, Sydney Basin and South East Corner Bioregions
• Swamp sclerophyll forest on the coastal floodplains of the NSW North Coast, Sydney Basin and South East Corner Bioregions
• Subtropical Coastal floodplain forest of the NSW North Coast Bioregion
• Umina Coastal Sandplain Woodland in the Sydney Basin Bioregion
• Blue Gum High Forest
• Western Sydney Dry Rainforest in the Sydney Basin Bioregion
• Sydney Turpentine-Ironbark Forest
• Milton Ulladulla Subtropical Rainforest in the Sydney Basin Bioregion

Lantana may prove to be a threat to additional Endangered Ecological Communities.

22. L. camara is "regarded as one of the worst weeds in Australia because of its invasiveness, potential for spread, and economic and environmental impacts" (CRC Weed Management 2003). It is one of the initial 20 Weeds of National Significance declared under the National Weeds Strategy, and a national Lantana Strategic Plan has been adopted (ARMCANZ ANZECC&FM 2001). It is recognised in most States and Territories of actual or potential occurrence as a serious weed of agriculture or the environment or both.

23. L. camara is a declared Noxious Weed under the NSW Noxious Weeds Act 1993 in most areas of eastern NSW, although in some areas only particular flower-colour variants have hitherto been declared. As from March 1, 2006, all areas with prior listings of any variant now have the species aggregate declared as Control Class 4, except in the local government control areas of Bega, Eurobodalla and Lord Howe Island (all Class 3). A statewide declaration of all Lantana taxa as Class 5 Noxious Weeds applies to ban sale and movement.

24. Some 30 biological control agents for L. camara (29 insects and one rust) have been released in Australia since 1914 (Day et al. 2003; Walton 2004), with variable results that lack overall quantification but are assessed as "generally inadequate to stop spread" (Grice 2005). Research into new biological controls and integrated control methods is continuing.

25.The Invasion, establishment and spread of Lantana (Lantana camara L. sens. lat.) is eligible to be listed as a key threatening process as, in the opinion of the

Scientific Committee:

(a) it adversely affects threatened species, populations or ecological communities, or
(b) could cause species, populations or ecological communities that are not threatened to become threatened.

Associate Professor Lesley Hughes
Chairperson
Scientific Committee

Proposed Gazettal date: 08/09/06
Exhibition period: 08/09/06 - 03/11/06

References:

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