2 Protection Quarterly Vol.22(1) 2007 fl eshy tubers on both the (rhizomes to about 20 cm diameter) and at nodes on aerial stems. Aerial tubers are small irreg- ular ‘warty’ light brown or green in colour Review and variable in size from 5 mm to about 25 cm in diameter, often bearing numerous axillary buds. are subsessile, sub- cordate, cordate or with a petiole to 1–12 cm (and less frequently up to 15 cm) long, The biology of Australian weeds alternate, broadly ovate, or sometimes lan- ceolate, heart-shaped ± fl eshy to succulent 46. cordifolia (Ten.) Steenis depending on exposure; apex obtuse. The lamina is bright green in colour, darker A B C Gabrielle Vivian-Smith , Ben E. Lawson , Ian Turnbull and Paul O. green on upper surface, glossy, clammy DowneyD to touch, 1–15 cm long by 0.8–11 cm wide. A Queensland Department of Natural Resources, Mines and Water, Alan The infl orescence resembles a lamb’s tail, Fletcher Research Station, Sherwood, Queensland 4075, Australia. and is a raceme 6–65 cm long slender and Email: [email protected] drooping; either simple or 2–4 branched. B School of Natural and Rural Systems Management, The University of Flowers are about 3–5 mm in diameter, Queensland, St. Lucia, Queensland 4067, Australia. fragrant, greenish-white to cream-white, C numerous and short-lived. The corolla is Bellingen Shire Council, PO Box 117, Bellingen, New South Wales 2454, white, infl exed, the lobes ovate-oblong to Australia. elliptic, 1–3 mm long; apex blunt. Style D Pest Management Unit, Parks and Wildlife Division, Department of and stamens are white. The style is shorter Environment and Conservation (NSW), Hurstville, New South Wales 2220, than stamens, 3-cleft branching halfway Australia. along the fused part to form three stigmatic arms, each with a club- or clavate-shaped stigma. Filaments are narrow-triangular, widely divergent, bending outwards near Name base pedicels 1.5–3 mm long. Each fl ower The genus Anredera Juss. (family Basel- pinkish-red and herbaceous, then becom- is subtended by a minute persistent bract laceae), named after the Spanish word ing brown, exfoliating and woody with 1.5–1.8 mm long; receptacle is cup-shaped for or climber, enredadera, includes age and reaching 2–3 cm in diameter by two persistent hyaline bracteoles, the up to 12 of perennial, twining or (Sperling 1987). Madeira vine produces upper bracteoles greenish white, fl attened, scandent, succulent or mucilaginous, of- ten tuberous (Sperling 1987). Mem- bers of the genus are broadly distributed from southern USA (Florida and Texas) and Mexico, through Central and South (b) America, from Colombia to Argentina, as well as Cuba, the Caribbean and the Gala- pagos. (Ten.) Steenis (syn. Boussingaultia cordifolia Ten., B. basel- loides (non H.B.K.) Hook, B. gracilis Miers, (a) B. gracilis form pseudobaselloides Hauman, B. gracilis var. pseudobaselloides (Hauman) Bailey, Anredera baselloides Baill. and B. cor- data Spreng.) is native to South America, Paraguay to southern Brazil and north- ern Argentina (Wagner et al. 1999). The many synonyms are indicative of earlier confusion regarding the of the , as noted by Sperling (1987). The specifi c epithet, cordifolia, refers to the heart-shaped leaves of the species. Common names include Madeira vine, potato vine, lamb’s tail vine, mignonette vine, sweet mignonette, heart- Ma- deira vine, jalap vine, white shroud, en- (c) redadera del mosquito, enredadera papa (hereafter described by its main common name Madeira vine).

Description Madeira vine is a perennial, vigorous climbing vine, or in the absence of sup- Figure 1. The form and identifying features of Madeira vine with insets port, a scrambling shrub/ground cover. showing fl owers (a), three-cleft stigma (b), and shoots growing from an Stems are slender, twining and hair- aerial tuber (c) (drawing by Wilma Roux, reprinted from Henderson (2001) less to about 30 m long, initially green- with permission). Plant Protection Quarterly Vol.22(1) 2007 3 broadly elliptic to suborbicular, about 1–2 particular Argentina, Bolivia, Brazil, Para- eastern Queensland and north-eastern mm long and shorter than the perianth. guay and Uruguay (Xifreda et al. 1999). NSW. Crude estimates from this model Bracteoles are persistent after corolla is Starr et al. (2003) reported the species as suggest that 80 600 km2 (1%) of Australia is shed. The single seeded fruits are globose, ‘native from Paraguay to southern Brazil highly suitable, 278 000 km2 (4%) is suita- slightly compressed to triangular, 0.9–1.1 and northern Argentina’ [p.1], located ble, and 1 387 700 km2 (18%) is of marginal mm long crowned by the enlarged fl eshy roughly between 20 to 30°S latitude. Sper- suitability for growth of Madeira vine. tri-lobed style base, and loosely enclosed ling (1987), in a revision of the Basellaceae, in the fl oral cup (van Steenis 1957, Sper- notes the species is most abundant in its Habitat ling 1987, Webb et al. 1988, Harden 2000, native range in southern South America In its native range in South America, the Shu 2003). production is rarely ob- from sea level to 3000 m. Lawson (1999) species is located at 20 to 30°S latitude, served outside the native range (Sperling located records of Madeira vine on all with average temperatures ranging be- 1987, Swarbrick 1999). continents except Antarctica, includ- tween 20 to 30°C in January, and 10 to 30°C ing South America (Bolivia, Ecuador, in July. The region has an average annual History Paraguay, Peru), Mesoamerica (Costa Rica, rainfall of 500–2000 mm, comprising a di- In Australia, Madeira vine has naturalized Honduras, El Salvador), North America versity of vegetation types including for- following its introduction for ornamental (the southern United States), Asia (China), est, grassland, cropland, woodland and purposes (see Floyd 1989). Madeira vine Europe (France), Africa (Malawi, Senegal) scrub (Hammond 1986, cited in Starr et was a rare listing in early nursery cata- and Australia. It has also been recorded al. 2003). In New South Wales, the species logues from 1906 until 1920, with the fi rst on many islands throughout the Pacifi c is considered most problematic in coastal identifi ed listing being in a 1906 Melbourne region, including American Samoa, Cook areas with summer rainfall (Weeds Aus- nursery catalogue produced by George Islands, Kermadec Islands, Hawaii, Lord tralia 2005). Brunning and Sons in St. Kilda (Mulvaney Howe Island, Micronesia, New Zealand Many sources note localized incursions 1991). However, the catalogues examined and Norfolk Island. Additional records mainly in areas of human settlement, on by Mulvaney (1991) were predominantly show the species growing in Argentina, rainforest margins in coastal districts and from temperate areas (Sydney, Melbourne, Brazil, Italy, Malaysia, Uruguay (Xifreda riparian areas (e.g. Stanley and Ross 1983, Canberra and Adelaide), and Madeira vine et al. 1999), South Africa (Henderson 2001), Dunphy 1991, Harden 2000, Harden et al. may have been sold more commonly in Sri Lanka (Ranamukhaarachchi et al. 1997, 2004). Madeira vine is well recognized as tropical and subtropical regions where it Marambe 2000) and Thailand (Royal Forest a weed of urban gardens and waste areas is more frequently cultivated. Irrespective, Department of Thailand 2005). Although (Esler 1988, Blood 2001). In Western Aus- these records suggest a possible introduc- regarded as a tropical and sub-tropical tralia it has naturalized on the fringes of tion to Australia over 100 years ago. Her- species, the worldwide distribution of the urban lakes (Swarbrick and Skarratt 1994) barium records from Queensland suggest species suggests that its range extends into and has been found on Garden Island and that Madeira vine has been naturalized mild temperate climates. along creeklines on the Swan Coastal Plain for at least 55 years (Batianoff and But- Madeira vine is recorded in all Austral- (Plant Protection Society of Western Aus- ler 2002). The fi rst report of Madeira vine ian States and Territories, with the excep- tralia 2003). The species is a major problem being naturalized in New South Wales tion of the Northern Territory (Blood 2001) in lowland rainforest remnants and fertile occurred a few years later in the 1960s at (Figure 2). Records of Madeira vine extend fl oodplains (Floyd 1985, Anon. 2005) and Wingham Brush in the north east of the along the eastern seaboard of the continent in moist eucalypt forests (Swarbrick and State (Stockard 1983). By the late 1970s, the between Cairns (17°S) and Hobart (43°S). Skarratt 1994, Blood 2001, Anon. 2005). On infestation of Madeira vine at Wingham The species has been recorded in Western moist fertile soils, particularly in the above Brush was such that mature trees had col- Australia around Perth (Csurhes and Ed- forest types, the species is an aggressive lapsed under the weight of the vines. By wards 1998). A record of the species from weed that can climb 40 m into tree cano- the 1980s it was deemed to be problematic Eulo in western Queensland (Queensland pies (Sydney Weeds Committee 2005). over large areas of New South Wales and Herbarium HERBRECS record) refl ects Aerial tubers of the species persist on the southern Queensland (see Floyd 1985), the current inland limit of the species in forest fl oor, often covered by leaf litter and specifi cally in riparian vegetation, the edg- Australia (approximately 800 km from the silt from fl ooding (Floyd 1985, Harden et es of rainforest, tall open forest and damp coast), although it is reportedly spreading al. 2004). The species exhibits poor growth sclerophyll forests (Csurhes and Edwards along watercourses in inland New South in low light conditions, but grows rapidly 1998). In south-eastern Queensland the Wales (NSW) (Weeds Australia 2005). in high light environments (Floyd 1985). species was ranked as fi fth in terms of in- However, most Australian records are The effects of Madeira vine are most pro- vasiveness and impact of 200 environmen- located in south-eastern Queensland and nounced on the edges of vegetation rem- tal weeds (Batianoff and Butler 2002). Ma- northern NSW, which is consistent with nants and in canopy gaps (Dunphy 1991), deira vine is now naturalized in all states statements in the literature (Stanley and where it rapidly climbs into the canopy, except the Northern Territory (Swarbrick Ross 1983, Dunphy 1991, Swarbrick 1999) and covers canopy trees (Floyd 1985, King and Skarratt 1994, Blood 2002). While oc- that highlight the invasiveness of the spe- 1988) (Figure 4). currences of the vine in Victoria, Tasmania cies in this region. In addition to the moist vegetation types (Blood 2002), South Australia and Western An assessment of the potential distribu- detailed above, Carr et al. (1992) note that Australia (Swarbrick and Skarratt 1994) tion of Madeira vine in Australia, based the species presents a threat to rock outcrop remain fairly isolated, it is expanding in all upon its distribution overseas using vegetation in Victoria, while Blood (2001) of these states. For example, in Tasmania, CLIMEX for Windows, Version 1.1a (Suth- and West (2002) describe infestations of where there were no records of naturaliza- erst et al. 1999), suggests the species has a the species on cliff faces in Victoria and tion prior to 1970 (Rozefelds et al. 1999), high potential for spread (Lawson 1999) New Zealand, respectively. These are con- Madeira vine is now reported as problem- (Figure 3). The model indicates that the sistent with ‘The Flora of Crete’ (Chilton atic in the north of the state (University of species poses a signifi cant threat to large and Turland 2004) that lists the species as Tasmania 2003). areas of coastal and sub-coastal Australia, occurring on ‘cliffs, rocks, rocky slopes’. with some potential for growth in every Madeira vine is also recorded as a problem Distribution State and Territory. Predictions show in dry coastal vegetation (Carr et al. 1992, Madeira vine is native to tropical and that the growth potential of the species is Blood 2001). The occasional occurrence of sub-tropical areas of South America, in highest in the subtropical zone of south- the species on frontal dune systems (Blood 4 Plant Protection Quarterly Vol.22(1) 2007 2001), mangroves (Weeds Australia 2005, Relationships with other species Cultivated relatives G. Vivian-Smith personal observation) and Facilitation Madeira vine can be confused with Ceylon coastal bluffs (West 2002) suggests some The invasive vine, cat’s claw creeper or Malabar spinach ( L. or B. ru- salt tolerance. The species is documented (Macfadyena unguis-cati (L.) A. Gentry) has bra L.), a related species within the Basel- as having some frost tolerance (Stockard et been hypothesized as a facilitator species laceae that is cultivated in Australia. This al. 1985). Blood (2001) attributes Madeira for invasion of rainforest remnants and species is grown as a green leaf vegetable vine’s capacity to tolerate short periods of fragments by Madeira vine (Floyd 1989). in gardens in south-eastern Queensland adverse conditions, such as drought, snow The increased light levels resulting from and has fl eshy leaves, rapid growth and a and frost, to the die-back of aboveground canopy collapse by cat’s claw creeper are climbing habit very similar to Madeira vine shoots during such periods. thought to create more favourable growth (G. Vivian-Smith personal observation). It conditions for Madeira vine (Floyd 1989). is also cultivated in South America, Africa and Asia, where the leaves are used as a vegetable and for medicinal purposes (van Steenis 1957). It produces fl eshy, purple- reddish ‘pseudoberries’ with purple juice that is used as a food colouring (van Stee- nis 1957). Another related species, in the Basellaceae, that is traditionally cultivated as a native tuber crop in the Andes is ul- luco, Ullucus tuberosus Loz. It is not known to be present in Australia, but has recently been reintroduced to New Zealand where it is being evaluated as a potential food crop (Busch et al. 2000).

Growth and development Madeira vine shows prolifi c growth in high light environments, with seasonal growth of up to 6 m reported (van Steenis 1957). Other growth rates reported from anecdotal observations range from 1 m per month (Floyd 1989), to more than 1 m per week during warmer months (Stockard et al. 1985). Cummings (1999) measured stem growth, recording a mean stem elon- gation of 101.6 cm from potted tubers over Figure 2. The recorded distribution of Madeira vine in Australia, detailing eight weeks in a summer glasshouse ex- the sources of those records. Mainland regions are based on IBRA periment, suggesting that growth rates are bioregions in Australia. likely to vary with factors such as growing conditions and plant resource reserves. Reports indicate that Madeira vine growth 120°0'0"E 130°0'0"E 140°0'0"E 150°0'0"E is suppressed under low light conditions, #LIMATE3UITABILITY compared to other invasive vines, such as 10°0'0"S 10°0'0"S 5NSUITABLE M. unguis-cati (Stockard 1983, Stockard et -ARGINAL al. 1985, Floyd 1985, 1989). 3UITABLE In addition to vertical growth, Madei- (IGHLYSUITABLE ra vine also produces large resource re- serves in the form of subterranean tubers and aerial tubers that initially form in the 20°0'0"S 20°0'0"S leaf axils. Subterranean tubers, found to depths of 1 m, are commonly reported to reach 20 cm in diameter (Muyt 2001). Aerial tubers frequently reach 10–20 cm in diameter within established infestations (G. Vivian-Smith personal observation), 30°0'0"S 30°0'0"S but can reach 30 cm in diameter (Floyd 1989). Tubers drop off the parent vine or are dislodged through disturbance, grow- ing into new plantlets or ‘tuberlings’ un- der favourable conditions. Germination is phanerocotylar (i.e. 40°0'0"S 40°0'0"S free of seed coat) and epigeal (Swarbrick

0 300 600 900 1200 1500 1999). Seedlings are rarely observed, but Kilometers are described as having fl eshy oval-ovate 120°0'0"E 130°0'0"E 140°0'0"E 150°0'0"E cotyledons, a succulent hypocotyl and dis- tinct juvenile leaves 5–10 mm in length, Figure 3. The predicted potential distribution of Madeira vine in Australia germinating from soil depths of several based on CLIMEX climate model with arbitrarily assigned growth classes millimetres (Swarbrick 1999) (Figure 5). (after Lawson 1999). Plant Protection Quarterly Vol.22(1) 2007 5 Reproduction data). It was estimated that Floral biology and phenology between 0.2–5.4% of the dried Flowers appear bisexual but are often func- fl owers present contained ger- tionally unisexual (Sperling 1987). No pol- minable (G. Vivian-Smith lination studies of Madeira vine have been unpublished data). Genetic identifi ed. However, fl owers are scented or pollination tests have not and produced in a large display, with been conducted to determine stamens bearing a nectary at their base why these populations pro- (Sperling 1987), suggesting pollination by duce seed, while the majority insects. The fl owers of Madeira vine of populations in Australia are in Maui (Hawaii) were noted to attract po- reportedly sterile. As flower tential pollinators, including various Hy- production is prolifi c, seed in- menoptera (including bees, wasps, and put to the environment could ants) but did not set fruit (Starr et al. 2003). well be signifi cant, when it oc- Infl orescences of several other members of curs. Seeds germinated readily the genus are reported to be attractive to under moist, light conditions in fl ies in their native range (Sperling 1987). Petri dishes placed in a growth Flowers are present in summer, peaking in cabinet with an alternating 12 h autumn (in Australia, December to May) thermo/photoperiod (15/25°C) (Muyt 2001). (G. Vivian-Smith unpublished data). Seed production and dispersal The seed is contained in a Sperling (1987), in a revision of the Basel- globose, indehiscent capsule laceae, reported that seeds were rarely (0.9–1.1 mm long) that retains produced in cultivated plants, but were the perianth (Sperling 1987, Xi- not infrequent in wild plants from south- freda et al. 1999), and is without ern South America, with fruit production an obvious means of dispersal, common in Bolivia, Argentina, Uruguay, although Swarbrick (1999) spec- Paraguay and Brazil. Xifreda et al. (1999) ulates that movement of soil or noted that the occurrence of fruit with water may result in dispersal. seed production was one of several char- The retained perianth makes it acters, in addition to chromosome number, diffi cult to differentiate seeds Figure 4. An infestation of Madeira vine, that separated two intraspecifi c taxa in A. from dry, spent fl owers on old showing its growth habit. cordifolia, namely Anredera cordifolia (Ten.) infl orescences (potentially lim- Steen. subsp. cordifolia and A. cordifolia iting observations of seed set), subsp. gracilis (Miers) Xifreda & Argimón. and may also aid in seed disper- One of these taxa, A. cordifolia subspecies sal by enhancing buoyancy in cordifolia (n = 36 chromosomes), was found wind or water (G. Vivian-Smith to be autotriploid, and is described as the personal observation). more extensively cultivated herb with prolifi c tuber production. This subspecies Vegetative reproduction produces abnormal pollen grains, and was Madeira vine’s major means of concluded to have virtual sterility. Analy- reproduction is through asexual sis of herbarium and live plant samples tubers on roots and stems (Sper- from wild populations of the other sub- ling 1987). In addition to tubers, species, A. cordifolia subsp. gracilis (n = 24 vegetative reproduction can oc- chromosomes) indicated the production of cur from stems and rhizomes viable pollen, smaller fl owers and fruit. (Brickell 1996). Axillary tubers Madeira vine is reported not to produce are produced at nodes and can Figure 5. Photograph of a Madeira vine seed in New Zealand (Esler 1988), China be highly variable in morphol- seedling emerging from attached remains of (Shu 2003) or Hawaii (Starr et al. 2003). ogy, with some bearing many In Australia, seed production is consid- warty or fi nger-like protuber- the persistent perianth. Seeds were collected ered uncommon, with only one scientifi c ances, while others are more at Welk’s Remnant, south-eastern Queensland record at Redwood Park, Toowoomba, globose in shape. These tubers and germinated at the Alan Fletcher Research Queensland (27°33”43.7’S, 151°59”50.3’E) may be found at densities of Station in 1998. (Swarbrick 1999). Additional anecdotal re- over 1500 m-2 on the forest fl oor, ports of seed production exist from Welk’s under heavy infestations of the Remnant and Dwyers Scrub Conservation vine (Stockard 1983, Stockard et al. 1985, cultivation for ornamental purposes, with Park, in the adjoining Lockyer Valley (T. Floyd 1985). In other areas, tuber produc- early transport of the vine from South Armstrong personal communication). tion is also heavy, with reports of 7.5 tonnes America thought to be via Spanish and Swarbrick (1999) suggested that seed pro- of tubers removed from two locations over Portuguese traders (Sperling 1987). Unin- duction may only occur in the most fa- a four year eradication attempt on Raoul tentional escape into neighbouring natural vourable years and does not appear a ma- Island, New Zealand (West 2002). areas and over longer distances has oc- jor means of reproduction. In June 2005, curred via the dumping of garden waste viable seeds, confi rmed via germination Dispersal containing tubers, rhizomes and other tests, were collected from three of four Madeira vine is primarily dispersed by viable plant fragments (Starr et al. 2003). populations sampled on the Toowoom- anthropogenic means, gravity and water In large infestations, aerial tubers drop to ba Range, including the Redwood Park (Dunphy 1991). Intentional human-medi- the ground at maturity, following physi- population (G. Vivian-Smith unpublished ated spread of Madeira vine occurs via cal disturbance (G. Vivian-Smith personal 6 Plant Protection Quarterly Vol.22(1) 2007 observation), when the parent plants expe- surface and within the canopy of the vine, (0–30%) conditions (Lowe 2000). Cum- rience stress (Prior and Armstrong 2001), and largely comprises aerial and subter- mings (1999) also reported 100% tuber or following manual control (Stockard ranean tubers. In such species, the shoot establishment rates under irrigated glass- 1993, P. Downey personal observation). growth from tubers can be superior to that house conditions. Shade treatments (0, Measurements of tuber dispersal (month- of seedlings due to the higher resource re- 50 and 90%) in the glasshouse were also ly assessment of tuber deposition in raised serves available to the shoot, but tubers found to signifi cantly reduce shoot emer- traps placed under vine canopies) over 24 are frequently more vulnerable to desic- gence and growth (i.e. length and leaf months at infestations (n >5) in south-east- cation than seeds, leading to potentially number) from the tubers (Lowe 2000). ern Queensland indicate continuous tuber higher mortality (Cousens and Mortimer deposition throughout the year (G. Vivian- 1995). Importance Smith unpublished data). Information or data enabling the de- Detrimental Secondary dispersal of such tubers may velopment of a model describing the In Australia, Madeira vine was ranked occur following fl ooding events or other population dynamics of Madeira vine are 41st out of the 71 invasive plants consid- disturbances (Pallin 2000, Vivian-Smith lacking. Ideally such data would describe ered during a national assessment for the and Panetta 2002). Localized dispersal via the seasonal rates of tuber production for Weeds of National Signifi cance (Thorp and sea currents is also suspected (West 2002), different plant- and tuber-size classes, the Lynch 2000), highlighting its current and but the salinity tolerance of tubers has not length of time before independent ramets potential status as one of Australia worst been tested. The potential for freshwater begin tuber production, rates of vegeta- environmental weeds. In other countries water dispersal has been investigated by tive expansion (e.g. ramet production and Madeira vine is also a problematic weed. testing buoyancy of tubers placed in fl oat- growth), tuber and shoot mortality rates, For example, in Hawaii (USA) the species ing cages in an experimental pond (Viv- the effects of different environmental vari- is a listed noxious weed (Starr et al. 2003), ian-Smith and Panetta 2002). Madeira vine ables (e.g. light and moisture) on growth in parts of New Zealand it is a declared tubers showed poor fl oating ability, with and resource allocation, and the role of pest species, and in South Africa it is also most tubers sinking in less than a day. density dependence on these parameters. a declared weed (Henderson 2001). However, two of the 375 tubers tested re- Madeira vine is a signifi cant environ- mained fl oating and viable for the entire Persistence mental weed in New South Wales, Queens- 30 days duration of the experiment. This Anecdotal reports indicate that aerial land and Victoria, and a garden escape in suggests that occasional long distance dis- tubers can persist for long periods rang- South Australia and Western Australia. persal of individual tubers by water is pos- ing from two (Landcare 1995), to five In New South Wales, the impact of exot- sible. A signifi cant proportion (33%) of tu- (Stockard 1993), to 15 years (Harden et al. ic vines, such as Madeira vine, has been bers survived 30 days in water, indicating 2004). Subterranean tubers are thought to acknowledged with a preliminary deter- that tuber viability after prolonged fl ood- persist for between 5–10 years, with re- mination for listing as a key threatening ing could be signifi cant. Despite low tu- ports of resprouting occurring for up to process under the NSW Threatened Species ber buoyancy, dispersal of Madeira vine is 10 years despite annual herbicide appli- Conservation Act 1995 (see NSW SC 2005a). likely during major fl ooding events, when cation (Muyt 2001). Tuber persistence un- In the Northern Territory, Madeira vine is rafts of dislodged vegetation containing der experimental conditions is limited (G. listed by the Northern Territory Parks and vines and tubers are transported down- Vivian-Smith unpublished data). Of 1100 Wildlife Conservation Amendment Act 1993 stream. An additional disturbance result- tubers tested (ranging in size from <0.5–5 as a Prohibited Entrant. In Queensland, ing in secondary dispersal of Madeira vine g fresh weight), no viability was reported Madeira vine has been declared a Class 3 tubers is the nest mound construction ac- after two years from a range of treatments pest plant under the Land Protection (Pest tivities of brush turkeys (Alectura lathami including burial (0, 1 and 5 cm) or suspen- and Stock Route Management) Act 2002, Gray), potentially concentrating tubers in sion from an artifi cial canopy. making it illegal to introduce, supply or turkey nest mounds (Westermann 2000). sell the species anywhere in the state. In Lowe (2000) observed that vertebrate Establishment from tubers south-eastern Queensland, Madeira vine tuber herbivores could potentially act as Lowe (2000) tested establishment from was ranked 5th in terms of invasiveness dispersers via removal and partial con- tubers following a range of experimental and impact from a list of 1060 naturalized sumption of tubers. Via a simulated tuber disturbance treatments in several habitats plant species (Batianoff and Butler 2002). herbivory (cutting tubers in half) experi- (i.e. closed forest, open eucalypt forest and The NSW North Coast Weeds Advisory ment in the glasshouse, he found that her- cleared grassland). He found that shoot Committee ranked Madeira vine as the bivory treatments did not signifi cantly af- emergence was greater for larger (>10 g) 7th most invasive environmental weed fect shoot emergence or growth rates. than for smaller (0.5–10 g) tubers, but did in the region due to its impact and diffi - not vary with habitat types. Disturbance culty to control (NCWAC 2001). Madeira Hybrids treatments only altered emergence signifi - vine has also been listed as one of the 10 No records of hybridization in Madeira cantly in the most severe treatment (i.e. most serious invasive garden plants spe- vine were identifi ed. The work of Xifreda soil replacement with commercial potting cies currently available for sale by nurs- et al. (1999) (see earlier) suggests that the mix), while a treatment of surface vegeta- eries in New South Wales (Groves et al. virtual sterility of A. cordifolia subsp. cordi- tion and litter removal had no signifi cant 2005). Such prioritization has not yet re- folia would provide very limited opportu- effect. Shoot growth (changes in shoot sulted in listings under the NSW Noxious nity for hybridization in this subspecies. length and leaf number) from tubers was Weeds Act 1993 in the worst affected areas. not signifi cantly different between habi- Current declarations (13 in total) are all Population dynamics tats or disturbance treatments. Greater within the Sydney catchment (Campbell- There are no published studies of popula- tuber predation was observed in the two town, Hornsby, Hunters Hill, Ku-ring- tion dynamics of Madeira vine. Records forest habitats, particularly for the distur- gai, Lane Cove, Manly, Mosman, North indicate that in most cases, Madeira vine bance treatment involving soil replace- Sydney, Parramatta, Pittwater, Ryde, War- does not possess a seed bank, but has what ment (Lowe 2000). ringah, and Willoughby) and Lord Howe is referred to as a ‘bud bank’ or ‘meris- Shoot emergence from tubers in fi eld Island. At time of writing, the declaration tem bank’ (Cousens and Mortimer 1995). and glasshouse experiments indicated category W4c of the NSW Noxious Weeds The meristem bank of Madeira vine is mean shoot emergence rates were greater Act 1993 requires that: ‘The weed must contained below the ground, on the soil under glasshouse (45–100%) than fi eld not be sold, propagated or knowingly Plant Protection Quarterly Vol.22(1) 2007 7 distributed and the weed must be prevent- also notes that fi eld cases of poisoning Herbicides ed from spreading to an adjoining prop- attributed to Madeira vine are rare, com- The control of Madeira vine can be erty’. Madeira vine is considered most prising suspected poisoning of drinking achieved through the use of selective or problematic in south-eastern Queensland water by Madeira vine followed by sud- non-selective herbicides (Muyt 2001). (Stanley and Ross 1983) and northern New den death. In feeding trials, cattle fed 9.5 Challenges reported include the produc- South Wales (Dunphy 1991). It is consid- kg of freshly cut fl owering foliage exhib- tion of viscous exudates from cut stems ered the most destructive weed of north- ited no ill effects, while sheep and pigs fed which inhibit herbicide uptake (Stockard ern New South Wales rainforest remnants 1.8 kg over two days exhibited temporary 1993) and the diffi culty in achieving up- (King 1988, Dunphy 1991). Invasion by diarrhoea before recovering (Hurst 1942). ward translocation of herbicide and death Madeira vine is well documented for the Tuber toxicity appears to be untested. of the aerial tubers (Muyt 2001). alluvial rainforest remnants at Wingham Lowe (2000) noted tuber predation in fi eld Historically, bush regeneration meth- Brush (Stockard et al. 1985, Harden et al. plots by red-necked wallabies (Macropus odology has been applied to the manage- 2004) and Bellingen Island in the mid- rufogriseus Desmarest), rodents, slugs and ment of Madeira vine in Australia. Meth- north of New South Wales (Floyd 1989), snails. ods are varied, but frequently utilize the and a dry rainforest remnant at Rotary stem-scrape and paint, or stem-cut and Park in Lismore, northern New South Benefi cial paint methodology (Muyt 2001), or stem Wales (King 1988), all sites being the sub- Madeira vine has been introduced to many injection techniques (Stockard 1993). Typi- ject of intensive restoration efforts. areas of the world as an ornamental plant cally, the scrape and paint technique in- Madeira vine infestations threaten the for its thick, glossy foliage and tassels of volves carefully scraping a 20 cm length following endangered ecological com- fragrant white fl owers (Bailey 1975, Floyd of stem to expose the cambium and im- munities listed under the NSW Threatened 1989). Future Foods (2004) indicate that mediate application of undiluted herbi- Species Conservation Act 1995 – i) ‘Swamp both tubers and leaves, although unap- cide (e.g. Roundup® glyphosate 360 g L-1). Sclerophyll Forest on Coastal Floodplains pealing in taste and texture, can be eaten Cut and paint involves cutting the stem of the NSW North Coast, Sydney Basin raw or cooked; the authors note that no and immediately applying undiluted her- and South East Corner bioregions’ (See reports on edibility or potential toxic ef- bicide to the cut surfaces. Stem injection Keith and Scott, in press), ii) ‘Littoral fects were identifi ed. We suggest that the involves injecting vines with a pressurized rainforest in the NSW North Coast, Syd- investigation of potential toxicological syringe with 2–3 mL of Roundup to al- ney Basin and South East Corner biore- effects of Madeira vine tuber consump- low for translocation into the tubers. Basal gions’ (NSW SC 2004), and iii) ‘River-fl at tion by humans may be warranted, given bark applications of Vigilant® gel (piclo- eucalypt forest on coastal floodplains reports of toxic effects of consumption of ram 43 g L-1) to vines in Bellingen (New of the NSW North Coast, Sydney Basin Madeira vine on livestock reported in the South Wales) were observed to result in and South East Corner bioregions’ (NSW literature (see above). Madeira vine was successful control; however impact on aer- SC 2005b, Coutts-Smith and Downey, in often planted near outdoor toilets because ial tubers was not determined (I. Turnbull press). In addition, Madeira vine is also it was thought that eating the leaves could personal observation). considered detrimental to riparian vegeta- have a laxative effect (Anon. n.d.). The ax- Foliar applications are used where large tion, tall open forest and damp sclerophyll illary bulbils, leaves and tubers are used numbers of vines render the application forests (Floyd 1989). medicinally (Shu 2003). In Taiwan, the of the more labour-intensive, localized In the worst affected areas, the species species is used for its medicinal properties, application methods impractical. Gener- quickly covers the canopy, reducing trees with hypoglycaemic activity of extracts ally, vines are cut and allowed to reshoot to ‘vine-shrouded pole-like structures’ demonstrated in mice (Lin et al. 1988), and prior to herbicide application. Regrowth (Floyd 1989). Madeira vine is considered to anti-infl ammatory, liver-protective (Lin et from vines and sprouting tuberlings is be the heaviest of the problem vine species al. 1994), and relaxant effects of extracts then sprayed with 1:50 glyphosate:water in subtropical Australia, due to its prolifi c demonstrated in rats (Lin et al. 1997). The (Stockard 1993). This method has been tuber production and vegetative growth plant has also been used as a folk medicine found to control 70–80% of stem tubers (Floyd 1989). This often leads to collapse analgesic and for the symptomatic treat- (BRAIN 1995). Hayley (1997) cited in Starr of supporting vegetation (Stockard et al. ment of diabetes mellitus in Taiwan (Lin et et al. (2003) recommends, after all tubers 1985), further compromising the structure al. 1988). Ethnobotanical reports note use are removed, a foliar spray of Escort® and functioning of the affected vegetation. of Madeira vine for treatment of sexually (metsulfuron methyl 60% by weight), Germination and regeneration of native transmitted diseases (Tshikalange et al. Roundup (glyphosate 360 g L-1) and Pulse® species is inhibited by dense growth of 2005). The close relative of Madeira vine, (organosilicate penetrant/surfactant) on Madeira vine (Harden et al. 2004, Anon. A. versicaria (Lam.) C.F.Gaertn., is also cul- plants and tubers as soon as green sprouts 2005). tivated and used medicinally by wrapping have two or four leaves on each sprout. Madeira vine is also known to threaten the around bruises, frac- Timing of follow-up spraying is important four species listed as endangered under tures and broken bones (Sperling 1987). because if left too long, new subterranean the NSW Threatened Species Conservation tubers will form, prolonging the control Act 1995, being Illawarra socketwood Weed management effort. Wildy (2004) suggests applying a (Daphnandra sp. C. Illawarra (R. Schodde In Australia, Madeira vine has been recog- foliar spray of Garlon 4® (triclopyr) mixed 3475)) and ripple-leaf muttonwood (Ra- nized as a potential environmental weed with water at 5 mL L-1. Vine weed control panea sp. A Richmond River (J.H. Maid- and as a candidate species for preventative trials held at Murphy’s Creek in Queens- en & J.L. Boorman NSW 26751)), Nilsen control (Csurhes and Edwards 1998). Ef- land found that both fl uroxypyr and met- Park she-oak (Allocasuarina portuensis fective eradication or control of this plant sulfuron methyl high volume foliar appli- L.A.S.Johnson), and Coxens double-eyed takes considerable effort and patience cation killed Madeira vine, but regrowth fig parrot (Cyclopsitta diophthalma cox- over a long period of time (Blood 2002). from tubers occurred with both chemicals eni Gould) (Coutts-Smith and Downey in The size and longevity of subterranean tu- (Armstrong and Keegan 1996). Prior and press). bers, the number of aerial tubers produced Armstrong (2001) compared the effi cacy and the lack of susceptibility of the tubers of one, two and three foliar applications of Toxicity—Stanley and Ross (1983) note to herbicide present considerable manage- various concentrations of glyphosate and that Madeira vine can be poisonous to ment challenges (Muyt 2001). fl uroxypyr ester on Madeira vine. Flurox- stock but it is rarely eaten. Everist (1974) ypyr applied at 1 and 2 g L-1 of water and 8 Plant Protection Quarterly Vol.22(1) 2007 glyphosate applied at 3.6 and 7.2 g L-1 of have collected several potential biocontrol References water were effective in controlling treat- agents for potential use in South Africa Anon. (n.d.). Anredera cordifolia. www. ed stems. However, fl uroxypyr at these (Gandolfo et al. 2004, van der Westhuizen mullum.com.au/wilsoncreeklandcare/ rates was the only herbicide resulting in 2005, 2006). These surveys have resulted weeds.maderia_vine.html). signifi cantly less regrowth of new stems in the identifi cation of damaging insects, Anon. (2005). Weeds of the Blue Moun- from subterranean tubers in the months including leaf-feeding chrysomelids and tains bushland. Garden plants going between applications. These authors also lepidopterans, as well as a fungal patho- wild – a guide to identifi cation and reported that the removal of competing gen (van der Westhuizen 2005). One po- control. www.weedsbluemountains. vegetation through the use of the non-se- tential candidate, a leaf-feeder Plectonycha org.au/madeira_vine.asp. Accessed 15 lective herbicide glyphosate may favour correntina Lac. (Chrysomelidae), is un- May 2005. regrowth from subterranean tubers, es- dergoing host-specifi city testing in South Armstrong, T. and Keegan, S. (1996). Ob- pecially if applied when translocation of Africa. Results indicate that this species servations on methods of controlling herbicides to the roots is low. has a narrow host range, with feeding and weeds at Murphy’s Creek. Proceed- The Australian Pesticide and Veterinary development occurring on some related ings of the 4th Queensland Weeds Medicines Authority (APVMA) database species, including Basella alba, Ullucus Symposium, Longreach, pp. 86-9. Weed reports only three products with Madeira tuberosus (both Basellaceae) and Talinum Science Society of Queensland, Bris- vine included on their label; all contain paniculatum (Jacq.) Gaertn. (Portulacaceae) bane. fl uroxypyr (220 g L-1) as the active ingre- (Gandolfo et al. 2004). The authors suggest Armstrong, T. and Prior, S. (1997). Con- dient. Two Minor Use permits (7485 and that this agent has potential for biocontrol. trol of Madeira vine (Anredera cordifolia) 5206) allow for the use of glyphosate at 10 A damaging leaf spot fungus, Alternaria using mechanical methods, herbicides g L-1 and 1:50 respectively (APVMA 2005). alternata Fr.: Fr. Keissl., has also been re- and hot fi re. In ‘Alan Fletcher Research ported on Madeira vine in Taiwan (Lai et Station: Environmental weed manage- Mechanical al. 1996), but has not been considered for ment research’, pp. 37-43. (Queensland Mechanical removal is generally only con- biocontrol. Department of Natural Resources, Bris- sidered practical in localized infestations Madeira vine has also been considered bane). where follow-up control, either mechani- as a target for biological control in New APVMA (2005). Australian Pesticide and cal or chemical, is readily available. Sub- Zealand (Syrett 2002). However, a ranking Veterinary Medicines Authority. www. terranean tubers can be dug out, but all tu- process for biological control of weed tar- apvma.gov.au/. Accessed 19 July bers must be removed to prevent regrowth get species in New Zealand downgraded 2005. (Muyt 2001). Vine cutting can result in in- the importance of Madeira vine due to its Bailey, L.H. (1975). ‘Manual of cultivated creased recruitment of plants from aerial lack of seed production (Syrett 2002). plants most commonly grown in the tubers falling from the dying vine (Stock- continental United States and Canada’. ard 1993). It is therefore recommended Cultural control (Macmillan Publishing Co., New York, that aerial tubers be removed from the The Bushland Friendly Nursery Scheme USA). vine where possible, prior to vine cutting operating on the NSW North Coast pro- Batianoff, G.N. and Butler, D.W. (2002). (Muyt 2001), with a tarpaulin being placed motes nurseries that do not sell environ- Assessment of invasive naturalized below the vine to facilitate collection and mental weeds, including Madeira vine plants in south-east Queensland. Plant disposal of falling tubers (BRAIN 1995, (BFNS 2005). Protection Quarterly 17, 27-34. Sutherland Shire 2003). Stockard (1993) BFNS (2005). Bushland Friendly Nursery reports that aerial tubers and stems held Fire Scheme. www.northcoastweeds.org.au. in the canopy can be pulled down using Trials undertaken by Armstrong and Prior Accessed 10 May 2005. aluminium extension poles with hooks (1997) found that the average growth of Blood, K. (2001). ‘Environmental weeds: attached; other workers recommend the aerial tubers was not signifi cantly reduced a fi eld guide for south-east Australia’. use of pruning saws to cut tubers from the following treatment with a Weed Drag- (C.H. Jerram and Associates – Science canopy (Muyt 2001). Disposal of collected on® propane burner. No published stud- Publishers, Mt. Waverley, Victoria). tubers is an important factor in preventing ies investigating the effects of prescribed Blood, K. (2002). Weed watch warning – reinfestation (West 2002). In a New Zea- autumn and spring fi res on Madeira vine Madeira vine, Anredera cordifolia. Under land eradication program on Raoul Island, infestations were identifi ed. Control 20, 10-1. more than 7.5 tonnes of tubers were col- BRAIN (1995). Brisbane Rainforest Ac- lected during a four year period, with dis- Acknowledgments tion and Information Network, ‘Weeds posal involving bagging tubers in plastic Moya Calvert, Marc Bryant (PestInfo Man- – Madeira vine’. www.brisrain.webcen- for transport, placing them in a desiccator, agement, Qld NRMW) and Craig Walton tral.com.au/newsletters/issue2/ma- and fi nally burning them (West 2002). (Qld Inter-agency Pest Management Com- deira.html. Accessed 10 May 2005. mittee) generously assisted with mapping. Brickell, C. (ed.) (1996). ‘The Royal Hor- Biological control David Yates (Botany Department, Univer- ticultural Society A–Z encyclopae- An application has been made to declare sity of Qld) is thanked for his advice and dia of garden plants’, p. 121. (Dorling Madeira vine a biological control target assistance. Lesley Henderson from PPRI Kindersley, London, UK). in Australia (Dhileepan and Palmer 2005), (South Africa) kindly granted permission Busch, J.M., Sangketkit, C., Savage, G.P., with the application currently under con- to reproduce Wilma Roux’s line drawing Martin, R.J., Halloy, S. and Deo, B. sideration by the Australian Weeds Com- of Madeira vine. Trevor Armstrong (Qld (2000). Nutritional analysis and sensory mittee (B. Wilson personal communica- NRMW) provided seedling photographs. evaluation of ulluco (Ullucus tuberosus tion). Madeira vine is considered to be a John Swarbrick, Trevor Armstrong and Ve- Loz) grown in New Zealand. Journal good biological control target in Australia, ronica Newberry (Toowoomba City Coun- of the Science of Food and Agriculture 80 with a high probability of fi nding an agent cil) assisted in locating Toowoomba popu- (15), 2232-40. with the required host-specifi city, as there lations. Tanya Grimshaw (Qld NRMW) Carr, G.W., Yugovic, J.V. and Robinson, are no native representatives of the family assisted with proof-reading. Shane Camp- K.E. (1992). ‘Environmental weed in- Basellaceae in Australia (Dhileepan and bell, Dane Panetta and an anonymous re- vasions in Victoria – conservation and Palmer 2005). Exploratory surveys for viewer provided valued input to an earlier management implications’. (Depart- natural enemies in Brazil and Argentina draft of this manuscript. ment of Conservation and Environment Plant Protection Quarterly Vol.22(1) 2007 9 and Ecological Horticulture Pty Ltd, %20Part1.pdf. Accessed 10 August Lowe, A. (2000). ‘The infl uence of habitat Melbourne). 2005. and disturbance on Madeira vine (An- Chilton, L. and Turland, N. (2004). Flora of Groves, R.H., Boden, R. and Lonsdale, redera cordifolia) tuber establishment’. Crete: Supplement II, Additions 1997- W.M. (2005). 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