212BOAW Asparagus.Indd

46 Plant Protection Quarterly Vol.21(2) 2006 the trnL-trnF region of the chloroplast ge- nome) obtained from several populations (L. Morin unpublished results). However, two populations of A. asparagoides, with Review the same morphology as the south-west- ern Cape form described by Kleinjan and Edwards (1999), were found at Donovan and Port McDonnell in south-eastern South Australia in July 2004 (K.L. Batch- elor and P.J. Turner unpublished results). The biology of Australian weeds ITS sequences of these populations exactly matched that of South African populations 44. Asparagus asparagoides (L.) Druce of the south-western Cape form (from Princess Vlei and Silvermine in the West- Louise MorinA,C,D, Kathryn L. Batchelor B,C and John K. Scott B,C,D ern Cape) (L. Morin unpublished results). A CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia. The sequence of the south-western Cape B CSIRO Entomology, Private Bag 5, PO Wembley, Western Australia 6913, form differed by 32 base pairs (5% differ- Australia. ence) from that of the widespread form of C Cooperative Research Centre for Weed Management Systems, 1995–2002. A. asparagoides found in Australia. Based D Cooperative Research Centre for Australian Weed Management. on recent surveys, this new form appears to be restricted to south-eastern South Australia and western Victoria (Coles et al. 2006). A molecular phylogeny of all Name Most recent scientifi c papers incorrectly southern African broad-leaved Asparagus Botanical name credit authorship of A. asparagoides to W. species is required to determine if the two Asparagus asparagoides (L.) Druce (syno- Wight, perpetuating the mistake made by forms of A. asparagoides warrant separa- nyms: Medeola asparagoides L., Myrsiphyl- Fellingham and Meyer (1995) upon return- tion at the species level. lum asparagoides (L.) Willd, Asparagus ing the genus Myrsiphyllum to Asparagus. Information provided hereafter refers medeoloides (L.f.) Thunb., Elide asparagoides Asparagus asparagoides (L.) Wight is illegiti- only to the widespread form of A. aspara- (L.) Kerguélen, Dracaena medeoloides L.f., mate as Wight (1909) mentions the name goides that occurs across southern Aus- Elachanthera sewelliae F.Muell., Luzuriaga under Myrsiphyllum without citing the tralia. sewelliae (F.Muell.) K.Krause) belongs to name that it was based on, Medeola aspara- the family Asparagaceae. The species in goides L., or mentioning authorship of the Common names this family were previously included in combination A. asparagoides (Anon. 2006a). Asparagus asparagoides is commonly re- the Liliaceae sensu lato (Dahlgren et al. Druce (1914) was the fi rst to legitimately ferred to as bridal creeper in Australia, 1985). publish the combination A. asparagoides most likely because its foliage was used Over the years, the higher taxonomy (Obermeyer 1984, 1992, Anon. 2005a). in fl oral arrangements, particularly bridal of the Asparagaceae has oscillated be- Two forms of A. asparagoides have re- bouquets (Scott 1995, Parsons and Cuth- tween one and three genera and varying cently been recognized in South Africa: a bertson 2001). It is also sometimes called numbers of sub-genera (Kleinjan and widespread form that corresponds to the smilax or fl orist’s smilax for its resem- Edwards 1999). Obermeyer’s (1984) revi- most common form of the weed that oc- blance to members of the Smilacaceae, sion of the family, which divided it into curs in Australia, and a form restricted to although it is not related to that family. three genera, Asparagus, Myrsiphyllum and the south-western Cape (Kleinjan and Ed- Bridal veil creeper is another name used Protasparagus was widely accepted until wards 1999). The latter can be differentiat- in some States; however use of that name recently. The latest Australian treatment ed from the widespread form by rhizomes can be confusing because it also refers to of the Asparagaceae (Clifford and Conran that grow primarily upwards towards the another asparagus weed, Asparagus dec- 1987) included A. asparagoides in the ge- soil surface, larger tubers and cladodes linatus L. (Keighery 1996, Lawrie 2004). nus Myrsiphyllum following Obermeyer’s (leaf-like stems) that are generally more Parsons and Cuthbertson (2001) also list (1984) revision. In a subsequent revision, waxy and elongated. However, above- gnarboola, narbas and krulkransie (S. Af- Malcomber and Demissew (1993) con- ground growth in A. asparagoides is highly rica) as common names. In the USA A. cluded that Asparagaceae contains only plastic and thus not a reliable factor for asparagoides is commonly referred to as one genus, Asparagus, with two subgenera, separation of the two forms (Kleinjan and African asparagus fern (Anon. 2005a). In Asparagus (which includes Obermeyer’s Edwards 1999). Seedlings of the two forms contrast, in Australia asparagus fern refers Protasparagus) and Myrsiphyllum. More re- are indistinguishable at the macroscopic to a range of asparagus weeds including cently, Fellingham and Meyer (1995) up- level. Obermeyer’s (1984) description of Asparagus africanus Lam., A. aethiopicus L. held the decision that the family should A. asparagoides is clearly that of the wide- and A. scandens Thunb. (Shepherd et al. comprise a single genus, but argued that spread form, but her distribution map is 2001). the subgeneric status of Asparagus and incorrect because it includes the south- Myrsiphyllum is not warranted. Fukuda et western Cape form as well as some other Description al. (2005) recently examined the molecu- broad-cladode Asparagus species (Kleinjan The following description of bridal creep- lar phylogeny of Asparagus as inferred and Edwards 1999). er (Figures 1, 2) was compiled mostly from from plastid petB intron and petD-rpoA The winter-rainfall region of the West- Obermeyer (1984), Clifford and Conran intergenic spacer sequences. They found ern Cape Province is likely to be the geo- (1987), Scott and Kleinjan (1991), Klein- evidence that southern African species graphic source of most founder popula- jan and Edwards (1999) and Parsons and are potentially paraphyletic, supporting tions of A. asparagoides in Australia, based Cuthbertson (2001). sub-division of Asparagus into more than on a comparison of DNA sequence data three groups. In this molecular phylogeny (non-coding internal transcribed spacer Habit A. asparagoides was the most different from regions of the nuclear ribosomal DNA Bridal creeper is a geophyte with exten- all other Asparagus species. [ITS] and intron and intergenic spacer of sive, perennial, below-ground storage Plant Protection Quarterly Vol.21(2) 2006 47 organs and above-ground parts that die back annually or when conditions are un- favourable (Humphries et al. 1991). It pro- duces thin, wiry, twisting shoots, slightly woody at the base and up to 6 m long when offered support. Shoots emerging from the below-ground root system entwine with each other and surrounding vegetation, al- lowing them to climb understorey shrubs and small trees (Figure 1).

Leaves The assimilating organs of bridal creeper are not true leaves but fl attened, leaf-like stems called cladodes or phylloclades, arising in the axils of reduced, scale-like leaves (Arber 1924) (Figures 2a,b). The cladodes are stalkless, broadly ovate to lanceolate but sharply pointed with a smooth or minutely denticulate margin, 10–70 mm long, 4–30 mm wide, dark glossy-green when growing in shade, but Figure 1. Bridal creeper climbing shrubs and trees in an infestation at dull and light green in exposed locations Yanchep National Park in Western Australia. (Photo CSIRO, L. Morin). and have a delicate parallel venation with no prominent mid-vein. They are solitary and alternate along the stem, or borne in (a) groups on short side branches.

Flowers Flowers are sweetly-scented, 8–9 mm wide and 5–6 mm long when fully expanded (Figure 2c). They are borne on 3–8 mm long and slightly bent pedicels, singly or in pairs in the axils of the reduced scale-like leaves. The six tepals (term used when pet- als and sepals are similar in appearance) are greenish-white, turned backward and (b) 5 cm (c) fused into a tube in the lower half. The six stamens are connivent (touching but not fused) and slightly shorter than the tepals. Flowers develop in late winter and early spring.

Fruits Bridal creeper produces globular berries, 6–10 mm wide, initially green and ripen- ing red (Figure 2d). Mature berries are 2–4 mm wide and sticky. They contain 0–4 and rarely up to 14 black, shiny, spherical or ovoid seeds (K.L. Batchelor and J.K. Scott 1 cm unpublished results). Fruits are produced in mid-spring, ripen in early summer and can be retained on senescing plants for Figure 2. (approximate scale several months. (d) included). (a) Line drawings of Below-ground biomass bridal creeper cladodes (leaf- The below-ground biomass of bridal like stems) arising in the axils of creeper consists of a cylindrical, slender reduced, scale-like leaves. From (about 5 mm wide), branching rhizome the Tamar Valley Weed Strategy (underground stem with shoot buds) (Anon. 2005b). (b) Bridal creeper growing parallel to the soil surface and foliage consisting of thin, wiry and bearing numerous, radially-arranged, twisting stems and cladodes (Photo spindle-shaped, fl eshy tubers, 25–42 mm CSIRO, J.K. Scott). (c) Bridal creeper long and 8–20 mm wide, often continuing as roots (Figure 3). Rhizomes and tubers fl owers are borne on pedicels arising of large plants entwine, forming a dense in the axils of leaves (Photo NSW mat about 5–10 cm thick just below the 1 cm Department of Primary Industries, J. soil surface. Hosking). (d) Bridal creeper mature red berries (Photo CSIRO, L. Morin). 48 Plant Protection Quarterly Vol.21(2) 2006

1 cm

1 cm

Figure 3. Below-ground biomass of bridal creeper (approximate scale included). (a) Mass of fl eshy tubers with roots spirally attached to a central rhizome. (b) Branching rhizome separated from its tubers. (Photos CSIRO, A.J. Willis and C. Davies).

History 1991). Localized infestations of bridal agents have not been released extensively Bridal creeper, native to South Africa, creeper have been present in South Aus- because of the bridal creeper eradication was introduced to Australia as a garden tralia since the 1950s, and bridal creeper campaign (Cooper and Warren 2006). It ornamental possibly via Europe, where it is now widespread across southern South also does not include infestations on Lord was fi rst used in horticulture (Jessop 1986, Australia, including Kangaroo Island and Howe Island. Scott 1995). The 1857 plant catalogue of the Riverland (Robertson 1983, Cooke Climate modelling to predict potential William MacArthur, who supplied seeds and Robertson 1990, Young 1995). Bridal distribution of bridal creeper showed that to New South Wales, Victoria and South creeper has also been reported through- slight northern expansions could be ex- Australia, represents the earliest known out much of Victoria, with heaviest infes- pected along the east and west coasts of record of bridal creeper being available tations found in coastal areas such as Port the mainland (Scott 1995, Pheloung and for sale in Australia (Mulvaney 1991). It Phillip Bay and inland in the Horsham Scott 1996, Scott and Batchelor 2006). was also listed in other nursery catalogues region, the north-east and along the Mur- Based on the model, bridal creeper also in the 1870s and 1880s (Mulvaney 1991, ray River (Beauglehole 1988, Conran 1994, has the potential to spread extensively on Brookes and Barley 1992) and recorded as Pritchard 2002). In south-western Western the north and east coasts of Tasmania. a plant growing in the Adelaide and Mel- Australia, widespread but localized infes- bourne Botanic Gardens in 1871 and 1883, tations were reported in the 1990s (Scott Other countries respectively (Scott 1995). and Pigott 1993, Pigott and Farrell 1996, The native range of bridal creeper is south- By the early 1900s, bridal creeper was Pigott and Lund 1996). ern Africa, but it is also recorded from Na- widely used by fl orists for decoration and In New South Wales, bridal creeper mibia and further north in tropical Africa as greenery in bridal bouquets (Scott 1995). was listed in 1981 as present on the central (Obermeyer 1984). Kleinjan and Edwards It was also extensively used as a garden coast, southern western slopes and western (1999), however, pointed out that the ve- ornamental at that time. Bridal creeper’s plains botanical regions and on Lord Howe racity of these northern herbarium records popularity with fl orists and gardeners was Island (Jacobs and Pickard 1981). Indeed, it cannot be ascertained due to the absence short-lived and declined steadily during was fi rst recorded on Lord Howe Island in of tubers on specimens and consequently the fi rst part of the twentieth century. It 1962 (Le Cussan 2006). Until the mid-1990s additional collections are required to con- was fi rst recorded as naturalized in Vic- bridal creeper was thought to be absent fi rm these records. toria in 1886 (Parsons and Cuthbertson from Tasmania, but several small infesta- Bridal creeper is not considered a weed 2001), although the fi rst herbarium speci- tions have since been reported. These, as in South Africa (Wells et al. 1986), although men of naturalized plants for that State well as infestations occurring on Flinders it is widely distributed and occurs in the was collected in 1943 (Clifford and Con- Island, are currently targeted for eradica- three main rainfall zones (winter, sum- ran 1987). The earliest herbarium records tion (Cooper and Warren 2006). Based on mer and even rainfall) (Kleinjan and Ed- of naturalized plants for South Australia herbarium records, bridal creeper has also wards 1999). It does not, however, occur and Western Australia were made in 1937 been found in the cooler areas of Queens- in the dry interior of the Western Cape and and 1960, respectively (Kloot 1986, Scott land in the south-eastern Darling Downs, Northern Cape Provinces or the hot sub- 1995). Warwick, Killarney and Toowoomba (R. J. tropical coast of KwaZulu-Natal. Henderson personal communication). Bridal creeper naturalized in New Zea- Distribution Bridal creeper has spread considerably land between 1900 and 1940 (Esler and Australia in the last two decades and is now widely Astridge 1987) and is now considered a Bridal creeper is naturalized in all States, distributed across southern Australia (Fig- weed in that country (Roy et al. 2004). It but not in the Northern Territory and the ure 4). The map of locations where bridal is also naturalized in some counties of Australian Capital Territory. Its initial dis- creeper biological control agents have California, USA, and although infestations tribution was clearly linked to locations been released, combined with herbarium are not large at this stage, managers have of early settlements (Carr 1996, Grant records from Australia’s Virtual Herbar- been alerted to its potential invasiveness 1996, Pigott and Farrell 1996, Stansbury ium (Anon. 2006b), is currently the most (Anon. 2005a). Climate modelling showed and Scott 1999). It is most prevalent in the accurate illustration of the national dis- that large areas of the Californian coast are temperate and Mediterranean regions of tribution of this weed. It underestimates, suitable for bridal creeper establishment southern Australia (Scott and Kleinjan however, distribution in Tasmania, where (Randall and Lloyd 2002). Bridal creeper is Plant Protection Quarterly Vol.21(2) 2006 49 as drainage lines (Pigott and Farrell 1996). It favours limestone outcrops in Yanchep National Park, Western Australia, which retain water and are well insulated from summer heat (Pike 1996). Stansbury (1999a,b) found that it grew best at sites with higher levels of available nitrates, po- tassium and iron. A recent study in south- western Western Australia has shown that soils where bridal creeper dominates have higher levels of available phosphorus than nearby weed-free areas (P.J. Turner per- sonal communication). It is unknown at this stage whether bridal creeper original- ly invaded phosphorus-rich areas or if it is modifying the soil environment. Bridal creeper has been recorded grow- ing in soils between pH 5.8 and 8.5 (Yates 1997, Stansbury 1999a,b, Raymond 1999, P.J. Turner and K. Siderov personal com- munications).

Plant associations Bridal creeper is found in a wide range of habitats including coastal heath on sandy Figure 4. National distribution of bridal creeper based on locations of sites dunes, eucalyptus and banksia woodlands where biological control agents have been released since 1999 (open circles; or forests, creek and river banks, swamps, based on information currently found in the CSIRO database; http://www. dry coastal vegetation, mallee shrubland, ento.csiro.au/weeds/bridalcreeper/project.html) with additional locations dry and damp sclerophyll open-forest, included based on herbarium records from Australia’s Virtual Herbarium and littoral rainforest (Cooke and Rob- ertson 1990, Scott and Kleinjan 1991, Carr (solid squares; Anon. 2006b). A herbarium specimen of Elachanthera 1996, Grant 1996, Pigott and Farrell 1996, sewelliae, a synonym of A. asparagoides, collected in 1886 at Nickol Bay in Yates 1997, Williams and Gerrand 1998, the Pilbara Region of northern Western Australia is not included in this Raymond 1999, Stansbury 1999a,b). Bridal map because bridal creeper was not found in that area during a survey in creeper-infested vegetation on roadsides the 1970s (Scott and Kleinjan 1991). and along tracks is believed to act as a source of infestation for surrounding ar- eas (Young 1995, Siderov and Ainsworth recorded as locally naturalized but not in- either in the winter, summer or aseason- 2004, Horlock 2005, Siderov et al. 2006). vasive in the Azores, Canary Islands, Por- ally (Kleinjan and Edwards 1999). Recent Bridal creeper is also one of the most seri- tugal and Sicily (Valdés 1980, Obermeyer South African surveys and examination ous weeds of citrus orchards in the Riv- 1984, Anon. 2006a). of herbarium specimens revealed that it erina, Sunraysia and Riverland irrigation cannot survive in arid or hot subtropical areas of New South Wales, Victoria and Habitat areas, and prefers the cool climate of high South Australia (Cooke and Robertson Bridal creeper is capable of invading a altitudes in north-eastern regions of South 1990, Kwong et al. 2002, Kwong and Hol- variety of habitats in warm temperate cli- Africa (Kleinjan and Edwards 1999). land-Clift 2004). However, it does not mates. In contrast to most other environ- In Western Australia, bridal creeper is persist in other agricultural ecosystems, mental weeds, bridal creeper’s invasion of associated with regions that receive more particularly pastures, as plants are read- new sites appears unrelated to disturbance than 350 mm of annual rainfall (Stansbury ily grazed (Siderov and Ainsworth 2004, events (Hobbs 1991, Raymond 1995, 1999, and Scott 1999). Cooke and Robertson Siderov et al. 2006). Bridal creeper invades Siderov et al. 2005). Stansbury and Scott (1990) reported a similar fi nding for South pine plantations, but it is not perceived to (1999), however, found via a questionnaire Australia, where bridal creeper was not have a signifi cant impact on tree growth (J. distributed to landholders that there was a recorded in areas with less than 300 mm Virtue personal communication). In South weak association between properties that annual rainfall. Bridal creeper can grow in Africa, bridal creeper mainly occurs as a had a long history of disturbance through areas where rainfall is sub-optimal provid- minor understorey species (Kleinjan and clearing of native vegetation and the pres- ing it is irrigated (e.g. private gardens or Edwards 1999). ence of bridal creeper. Bridal creeper pre- citrus orchards) (Young 1995), or in close fers shaded or part-shaded habitat (Meney proximity to watercourses or in moist gul- Growth and development et al. 2002). It does grow in hind dunes lies (Pigott and Farrell 1996, Stansbury and Morphology on exposed beaches in South Australia, Scott 1999). Bridal creeper seedlings have a single coastal cliffs and amongst shrubs closer to cotyledon and their above-ground parts shoreline in sheltered bays (J. Virtue per- Substratum develop slowly until the root system is es- sonal communication). Bridal creeper is able to grow on a wide tablished (Parsons and Cuthbertson 2001). range of soils, but prefers light-textured Cooke and Robertson (1990) recorded the Climatic requirements sandy and loam soils (Cooke and Robert- fi rst tuberous root at nine weeks after emer- Bridal creeper’s ability to alter its phenolo- son 1990, Grant 1996, Yates 1997, Stans- gence, and found that some seedlings were gy enables it to survive in different rainfall bury 1999a,b). It is particularly vigorous in capable of surviving their fi rst summer zones of South Africa, which receive be- soils with high moisture content (Meney et with this limited reserve. In Western Aus- tween 400 and 750 mm of annual rainfall al. 2002) and areas high in nutrients such tralia, seedlings produced an average of 50 Plant Protection Quarterly Vol.21(2) 2006 1.3–2.4 tubers over six months, depending �� on habitat (Stansbury 1999a,b). Robertson (1983) observed that young plants gener- ally had several stems and approximately � 20 tubers before a distinct short rhizome was produced. The rhizome develops into several branches comprising meristem- atic buds (growing points) and numerous, � fl eshy tuberous roots (Figure 3). Mature plants are characterized by their extensive below-ground mat of rhi- � zomes and tubers that make up more than 87% of the plant’s total biomass (Raymond 1995, 1996, 1999) (Figure 3). Raymond � (1995, 1996, 1999) recorded an average dry

weight of rhizome and tuber mats of 0.83 ������������������������������������ kg m-2 at one of her study sites in Victo- ria (corresponding to approximately 6000 �� �� �� �� �� �� tubers m-2), while at Western Australian sites the dry weight of rhizome and tuber �������������������������������� mat ranged between 2.3 and 3.7 kg m-2 (P.J. Turner personal communication). New shoots arise from meristematic buds on Figure 5. Correlation between above-ground and below-ground biomass of the rhizomes and not from tubers (Cooke bridal creeper. Reprinted from Biological Control, Vol. 30, Kleinjan, C.A., and Robertson 1990). Plants can be gen- Edwards, P.B. and Hoffmann, J.H., Impact of foliage feeding by Zygina erated from sections of rhizome as long sp. on tuber biomass and reproduction of Asparagus asparagoides (L.): as they bear at least one growing tip and relevance to biological control in Australia, pp. 36-41, Copyright 2004, with a few tubers (Kleinjan et al. 2004a). Indi- permission from Elsevier. vidual plants cannot be distinguished in dense infestations. Plants produce shoots early in the Physiology treatment (100% every fortnight for 20 growing season that can extend several No detailed studies have been carried out weeks) completely exhausted tuber re- metres by climbing over vegetation or on bridal creeper’s evapotranspiration serves of some plants and prevented re- other kinds of support. The climbing, in- and photosynthetic and translocation ca- growth after the last defoliation (L. Morin tertwining shoots can create columns or pacity. and A.J. Willis unpublished results). curtains of foliage, smothering surround- Bridal creeper’s preference for shaded Once plants establish extensive below- ing vegetation (Figure 1). Mature plants or part-shaded habitats is an indication ground reserves they can produce shoots typically fl ower only after several years of that it can photosynthesize effectively un- at the onset of cooler temperatures, even in vegetative growth (Robertson 1983). der low light intensities. Maximum growth the absence of available external moisture of stems has been recorded at low light in- (Kleinjan and Edwards 1999, Raymond Perennation tensities in laboratory conditions (Courtot 1999). This can be attributed to fleshy Perennation is achieved by the persistence 1965). Following rapid growth and estab- tubers retaining high moisture content of the extensive below-ground storage or- lishment of shoots at the beginning of the over several months. In the absence of gans and the production of adventitious growing season in late summer and early rain, foliage growth is largely restricted buds on rhizomes. With these characteris- autumn, bridal creeper diverts some of its to stems with small and slender cladodes, tics, bridal creeper fi ts perfectly the classi- photosynthates towards tuber production. but it returns to normal as soon as rainfall fi cation of a geophytic life form (Raunkiaer Raymond (1996, 1999) estimated that new resumes. 1934). Its storage reserves not only provide tubers account for a maximum of only 8% Daily growth rates of bridal creeper’s energy for shoot growth, but also buffer of the below-ground biomass each year. seedlings and above- and below-ground plants against adverse conditions, there- This, however, may be an underestimate biomass of established plants have not by ensuring persistence over many years because new tubers gradually change col- been measured. Nevertheless, the relative (Raymond 1996, 1999). In winter-domi- our during the season and become diffi - growth rate of above-ground biomass (dry nant rainfall areas, bridal creeper senesces cult to distinguish from previous years’ weight) of young plants has been shown, at the onset of warm weather in late spring tubers. in a glasshouse experiment, to follow the and relies on its below-ground biomass to The above-ground biomass of bridal same trend as that of other species (Turn- survive hot, dry summer months. creeper is positively correlated to its be- er 2003). Following initial exponential Bridal creeper is a long-lived perennial low-ground biomass (Turner 2003, Klein- growth, bridal creeper’s relative growth plant. Presence of dark and shrivelled tu- jan et al. 2004a) (Figure 5). Raymond (1999) rate reached a plateau and then declined bers indicates that an infestation is more noted that bridal creeper’s use of its be- as plants matured. In contrast, the relative than three years old (Siderov and Ains- low-ground reserves to support foliage growth rate of tuber biomass peaked in worth 2004). Raymond (1999) estimated growth is likely to be minimal due to the the middle of the experiment after bridal that the infestation at her main study site absence of any decrease in tuber biomass creeper had developed extensive foliage. was at least 15 to 20 years old, based on over her two year study. Bridal creeper’s Mature plants can tolerate fi re (Yates an approximate average increase of 18.3 g tuber reserves are not easily depleted, as 1997, Willis et al. 2003). Hot summer fi res dry mass m-2 of new tubers per year. The shown in experiments where bridal creep- can destroy a large proportion of tubers ly- tuber and rhizome mass has also been er was mechanically defoliated at regular ing near the soil surface, but generally fi re found to persist for over eight years after intervals (Raymond 1999, L. Morin and does not affect below-ground reserves lo- bridal creeper had been killed by glypho- A.J. Willis unpublished results). Only cated deeper in the soil (Carr 1996, France sate (Turner and Virtue 2006). the most severe and regular defoliation 1996). Below-ground biomass of well Plant Protection Quarterly Vol.21(2) 2006 51 established infestations is also rarely com- ��� pletely killed with one-off application of foliar herbicides, probably due to the high root to shoot ratio, which makes it diffi cult ��� to get suffi cient herbicide through the fo-

liage and into the root system (Pritchard ��

�� ��

2002). � � � �

Phenology � � � In South Africa, bridal creeper adapts its � �� �

phenology to different seasonal rainfall � patterns (Kleinjan and Edwards 1999). �� It behaves similarly in Australia, where its major distribution is in winter-rainfall dominant regions. In most areas, shoots �� emerge from the below-ground rhizome either slightly before or with the onset of the fi rst late-summer or autumn rains. For � example, Raymond (1996, 1999) found � � � � � � � � � � � � � � � � � that shoots began emerging in April 1992 at her intensively monitored study site ���� ���� ���� on the Mornington Peninsula (Figure 6). However, a major rainfall event in January 1993 triggered emergence earlier than usu- ����� al in the following growing season. Fur- Figure 6. Shoot density recorded every six weeks from April 1992 to thermore, the emergence of bridal creeper February 1994 in permanent plots at a site in the Mornington Peninsula shoots from the below-ground biomass has been shown to be linked with decreas- National Park, Victoria. No shoots were present six weeks prior to the fi rst ing temperatures in autumn in the absence assessment. Reproduced with permission from Raymond (1999). of substantial rains (Stansbury 1995, Ray- mond 1999). Seeds readily germinate throughout indicated that senescence is probably more Shoots that emerge early in the grow- autumn and winter if conditions and mi- associated with a drastic decrease in mois- ing season are the only ones that will pro- crohabitat are adequate. Seedlings have ture levels rather than with increasing tem- duce fl oral structures and fruits (Raymond a good chance of surviving over the dry peratures. Such phenology has been ob- 1996, 1999, Stansbury 1999b). Buds fl ower summer period providing they have had served in areas of South Africa where rain- and wither within two weeks and, by the time to develop some below-ground re- fall is evenly distributed throughout the third week swelling of the ovary is usu- serves (Cooke and Robertson 1990, Ray- year (Kleinjan and Edwards 1999). Similar ally visible (Raymond 1999). Up to 40% mond 1999). phenology is found in areas such as Tam- of fl owers set fruit at a study site in South In Australia, fl owering generally oc- worth in north-eastern New South Wales, Australia (Robertson 1983), while fruit set curs in late winter to early spring (Rob- where there is slightly more summer than ranged between 3.5 and 25% at sites on the ertson 1983, Raymond 1995, 1999), while winter rainfall (J. Hosking personal com- Mornington Peninsula where more than in South Africa fl owers are produced as munication). Although there is always 80% of buds had fl owered (Raymond 1995 early as July (Kleinjan and Edwards 1999). some above-ground growth present under 1996, 1999). Raymond (1999) explained Flowers occur only on shoots from well- these conditions, there is still an annual that such low fruit set was probably due established plants (Robertson 1983). Fruits turnover of shoots with most of them dy- to a combination of climatic or site-specifi c develop in the spring and mature into red ing during the summer months. factors such as high intraspecifi c competi- berries in late spring to late summer, de- tion and stress among fl owering shoots, pending on the region. For example, fruits Mycorrhiza extreme shading, herbivory on buds and ripened much earlier (October to Novem- Mycorrhizal associations between bridal flowers and mouldiness of developing ber) at sites in Western Australia with a creeper and any fungi have not been in- fruits leading to abortion. However, it Mediterranean-type climate than at south- vestigated. It is noteworthy that coloniza- may simply be explained by the fact that ern Victorian sites (December to January) tion by vesicular-arbuscular mycorrhizal fl owering shoots were not climbing on a (Stansbury 1996, Raymond 1999). fungi has been observed in secondary but support (see below). Above-ground biomass begins to se- not storage roots of cultivated asparagus, nesce in mid to late spring. The onset Asparagus offi cinalis L. (Wacker et al. 1990). Seed production and dispersal of senescence is earlier when plants are Mature bridal creeper fruits contain, on exposed to more direct sunlight. Shoots Reproduction average, two to three seeds (Raymond typically senesce from the apex down and Floral biology 1999, Stansbury 1999b). In rare cases, the take several weeks to die back completely Bridal creeper fl owers are bisexual and number of seeds per fruit is as high as 14 (Raymond 1999). By December senescence self-compatible. Cross-pollination has (K.L. Batchelor and J.K. Scott unpublished is usually complete in areas with distinct been demonstrated in hand-pollinated results). Fruit production is generally pro- winter rainfall patterns (Robertson 1983, fl owers, but the level of outcrossing ver- lifi c on climbing shoots but rare or lim- Stansbury 1995, Kleinjan and Edwards sus selfi ng still remains to be determined ited on long prostrate stems (Kleinjan and 1999). (Raymond 1995, 1999). Honeybees, Apis Edwards 1999, Stansbury 1999a,b). For Not all shoots senesced during the sum- mellifera L., visited bridal creeper fl owers example, at Western Australian sites the mer months at the study site of Raymond in large numbers during the day at one of mean number of fruits produced by brid- (1995, 1996, 1999) in 1992–93 (Figure 6) be- the study sites of Raymond (1995, 1999), al creeper climbing on a 1 m wide × 2 m cause it was wetter than usual. This fi nding collecting both pollen and nectar. tall trellis ranged between 527 and 3787, 52 Plant Protection Quarterly Vol.21(2) 2006 while plants provided with no support habitats has the potential to increase the averaging 24°C than at sites where aver- produced 0 to 5 fruits m-2 (C.D. Stansbury rate of invasion of remnant vegetation age temperature was above 27°C. unpublished results). patches at a local scale (Siderov and Ains- The effect of smoke and heat shock Frugivorous birds are recognized as worth 2004, Siderov et al. 2006). treatment (90°C for 10 min) on germina- important contributors to seed dispersal Bridal creeper seeds are also dispersed tion of bridal creeper seeds has been in- in bridal creeper (Raymond 1995, 1999, by other animal vectors. Seeds have been vestigated, but results of different trials Stansbury 1996, 1999b, 2001). Over her observed in rabbit (Oryctolagus cuniculus have been contradictory, highlighting the two year study, Raymond (1995, 1999) ob- L.) droppings in the vicinity of warrens need for further work in this area (Willis served that birds were the primary dis- and there is anecdotal evidence that foxes et al. 2003). Emergence of bridal creeper persal agents of bridal creeper, removing (Vulpes vulpes L.) are dispersal agents (Gra- seedlings within four weeks of a wildfi re 58–78% of fruits. Bridal creeper berries ham and Mitchell 1996, Raymond 1995, at Stokes National Park, Western Austral- have similar physical characteristics and 1999). Fox baiting has been suggested as ia, suggested that seeds can survive fi re nutritional value to bird-dispersed fruits a way to control long-distance spread of (France 1996). of native Australian species in wet sclero- bridal creeper (Agriculture and Resource Raymond (1999) demonstrated that phyll forests (Raymond 1999). Fruits are Management Council of Australia and most bridal creeper seeds were naturally retained on plants long after shoots have New Zealand [ARMCANZ] et al. 2001). buried within three months and that the senesced and their bright colour makes Mud containing seeds that adheres to seed bank is mostly transient, with only them attractive to frugivorous birds. At animals, vehicles and machinery is also <5% carryover of viable seeds between least 12 bird species, both native and ex- believed to contribute to dispersal (Par- years at Mornington Peninsula National otic, have been recorded feeding on bridal sons and Cuthbertson 2001), although this Park. She observed that seeds persisted creeper berries and potentially dispersing would be of relatively minor importance. longer on the soil surface than buried at seeds in southern Australia (see Table 1 Graham and Mitchell (1996) also observed 2 or 5 cm, with more than 55% of surface in Stansbury 2001). Bridal creeper fruits, situations where seeds dispersed down- seeds and none of buried seeds viable after however, may not necessarily be their pref- stream by fl ood events. 18 months (Figure 7). Most viability loss erence. For example, silvereyes (Zosterops was attributed to decay occurring within lateralis Latham) in Western Australia were Physiology of seeds and germination three months of burial. During that pe- seen switching from bridal creeper berries Seeds of bridal creeper have an after-rip- riod more than 50% of seeds at both burial to newly ripened fruits of a nearby com- ening requirement that ensures they do depths and only 2% of surface seeds had mon fi g (Ficus carica L.) tree growing in the not germinate too soon after they are pro- germinated (Figure 7). Surface seeds began wild (Stansbury 1996). In South Australia, duced, when conditions are likely not to germinating only after they were covered common starlings (Sturnus vulgaris L.) are be optimal for seedling growth (Robertson by litter. Raymond (1999) did not observe believed to be the primary agent respon- 1983). Raymond (1999) found that seeds any evidence of seed predators, such as sible for bridal creeper spread to remote stored for 2–6 months germinated twice ants, removing seeds. areas on the Nullabor Plain (P. Sheridan as fast as fresh seeds, but the total number �� � personal communication). On the other of seeds that germinated was lower after hand, silvereyes are the main dispersers in 8 to 12-months’ storage. Passage of seeds Western Australia, the irrigated Riverland through the gut of silvereyes was also ob- �� area of South Australia, Victoria and Lord served to reduce the after-ripening period Howe Island (Raymond 1999, Stansbury and enhance germination by 1.2 to 1.5 �� 2001, Le Cussan 2006, P. Sheridan per- times compared to unpassed seeds. Ray- ��������������� sonal communication). Ground-foraging mond (1999) commented that gut passage �� birds, such as common blackbirds (Turdus of seeds may have removed germination merula L.) and emus (Dromaius novaehollan- inhibitors in the seed coat or increased � diae Latham) also contribute to secondary permeability to gases, but it is unlikely to dispersal of fruits (Loyn and French 1991, have infl uenced penetration by water since Raymond 1999). unpassed seeds imbibe water readily. �� � Bird dispersal of seeds is highly sto- Mature seeds can germinate over a chastic and consequently dispersal is wide range of temperatures (10–30°C) in �� diffi cult to predict precisely (Siderov et both constant light or darkness, but germi-

al. 2005). The dispersal pattern is infl u- nation is optimal at lower temperatures of �� enced by the behaviour of different bird approximately 15–20°C (Robertson 1983, species and habitat structure, since many Fox 1984, Raymond 1999). Willis et al. birds fl y to perch-trees to consume fruits (2003), however, observed a strong inhibi- �� ��������������������� (Raymond 1999, Siderov and Ainsworth tory effect of light on germination when 2004). Stansbury’s (2001) diffusion model fl uctuating light and temperature regimes � showed that bridal creeper distribution at were used. Bridal creeper seeds can also � �� �� �� �� �� Bold Park in Western Australia was corre- germinate when exposed to water poten- ��������������������������� lated with the fl ight pattern of silvereyes, tials as low as -0.44 MPa, making them with a mean distance for seed dispersal of moderately tolerant of moisture stress Figure 7. Fate of bridal creeper 90.5 m. Occasional long-distance dispersal (Raymond 1999). Raymond (1999), how- seeds located on the soil surface is also possible and may signifi cantly in- ever, showed that seed viability was not af- (circles) or buried at depths of 2 cm crease rate of spread at a landscape scale. fected by exposure to high (30°C) and low (squares) and 5 cm (triangles) over Based on silvereyes’ gut passage rates and (5°C) temperatures, or low water poten- time at a site in the Mornington fl ight speed, the maximum potential seed tials (<-0.44 MPa). The effect of fl uctuating Peninsula National Park, Victoria. dispersal distance has been estimated to temperatures on seed germination in the be 12 km (Stansbury 2001). Nevertheless, fi eld has never been examined. Nonethe- (a) persistence of seeds (viable but gradual, short-distance bird dispersal of less, Stansbury (1999a,b) found that more ungerminated), (b) germination of bridal creeper seeds along roadsides and seedlings survived at sites in Western seeds in the fi eld. Reproduced with wildlife corridors that connect disparate Australia with summer soil temperatures permission from Raymond (1999). Plant Protection Quarterly Vol.21(2) 2006 53 In another experiment conducted in the ������ You Yangs Regional Park, Victoria, Ray- mond (1999) obtained similar germination in undisturbed sites and sites exposed to a ��� single small-scale soil disturbance. None- theless, seedlings established more readily ��� when they germinated in microsites un- der tree canopies rather than in the open. �� Seedling establishment was also promoted ��� by litter cover. � Vegetative reproduction Whilst seeds remain the most important

means for bridal creeper dispersal, es- ��������������� ��� tablished plants can also increase in size through spread of branching rhizomes ��� bearing meristematic buds. Stansbury and Scott (1999) estimated from a land- holder survey that patches of about 10 m2 ��� expanded radially by approximately 0.6 m year-1. This expansion rate however, is ��� most likely an overestimate considering ��� ��� ��� ��� ��� ��� ��� the rate of new tuber production per year recorded by Raymond (1999) (see above). ���� ���� Rhizome fragmentation has been used to increase numbers of plants in the nursery ����� trade (Bodkin 1986). Earthworks (e.g., roadside grading) can spread rhizomes Figure 8. Seedling dynamics over one year at a site in the Mornington over considerable distances to start new Peninsula National Park, Victoria; cumulative recruitment (circles), infestations. senesced seedlings (triangles) and total seedling number (squares). Reproduced with permission from Raymond (1999). Hybrids There are no known hybrids of bridal creeper. In South Africa, A. asparagoides is from the same location as those of the pre- 1999), and more studies are required to reported to grow sympatrically with oth- vious season. document variation between populations er closely-related species such as Aspara- Fruits are mostly produced on shoots across its distribution. gus ovatus T.M.Salter and A. kraussianus of established plants that emerge at the (Kunth) Macbride (Kleinjan and Edwards very beginning of the growing season Importance 1999), but no plants of indistinct (hybrid) (Raymond 1996, 1999). Fruit densities Detrimental form have ever been found at these sites have been recorded to be as high as 2149 Rapid growth of bridal creeper in autumn (P.B. Edwards personal communication). fruits m-2 in infestations where no artifi cial and its climbing habit provide canopy Artifi cial crosses between broad-leaved support was provided (Raymond 1999). dominance and make it highly competi- Asparagus species have not been attempt- Seedling recruitment is highly variable tive. Once a bridal creeper infestation is ed as far as we know. and fl uctuates from year to year and site established, the amount of light reaching to site. At one of Raymond’s (1999) sites on the soil surface is very low, thereby pre- Population dynamics the Mornington Peninsula, 80% of seed- venting other plants from persisting. Ray- At a local scale, vegetative extension lings emerged between April and June and mond (1999) reported that the amount of of below-ground rhizomes is the main reached an average of 214.5 seedlings m-2 irradiance reaching the soil surface was means of increase in area occupied by by August (Figure 8). More than 40% of reduced by 93% under a canopy of bridal bridal creeper. Local populations can also seedlings died during the growing season, creeper foliage. Competitiveness of bridal increase via spread of fragmented rhi- probably because of litter disturbance, and creeper is also due to the quantity of be- zomes. Average shoot densities up to 95.8 the remaining seedlings senesced at the low-ground reserves that occupy most of m-2 have been recorded at one study site in beginning of summer in December. The the space available in the substrate. Victoria (Raymond 1995, 1996, 1999) (Fig- following season up to 69% of the seed- Bridal creeper is recognized as one ure 6). At that site, shoots that emerged lings that survived the fi rst growing sea- of the most troublesome environmen- within one week of the commencement of son had re-emerged by June. Survivor- tal weeds in southern Australia, and is the growing season were more than three ship was likely affected by competition ranked in the top 20 Weeds of National times longer and produced signifi cantly for resources amongst seedlings and with Signifi cance (Thorp and Lynch 2000). It is a more cladodes than those that emerged established bridal creeper plants or other major problem for conservation because it in the following six weeks. Less than 20% species (Raymond 1999). can change the structure, fl oristic compo- of shoots died during the growing season, Life expectancy of bridal creeper can- sition and ecology of natural ecosystems. followed by high mortality at the onset of not be determined from established infes- A serious invader of both disturbed and the warm weather at the end of the season. tations because it is impossible to iden- undisturbed habitats, it can quickly domi- High levels of summer rainfall during the tify individual plants (Raymond 1999). nate understorey vegetation, modify the 1992-1993 summer in southern Victoria Long-term monitoring of seedlings to aesthetics, affect access and change the led to the survival of some shoots until reproductive maturity and beyond is re- overall structure of the landscape (Hobbs the following April, after new shoots had quired to assess longevity. There has only 1991, Humphries et al. 1991, Adair and started to emerge (Raymond 1995, 1996, been one study on the population dynam- Groves 1998, Raymond 1999, Siderov et 1999). These new shoots did not emerge ics of bridal creeper (Raymond 1995, 1996, al. 2005). 54 Plant Protection Quarterly Vol.21(2) 2006 Bridal creeper is a highly competitive also impede seedling establishment of na- orchards in Australia, where it smothers species that poses a direct threat to at least tive species by occupying all of the space trees, displaces citrus roots and reduces 16 native plant species in South Australia available in the substrate or by changing fruit production (Kwong et al. 2002, Kwong and New South Wales (Davies 1986, 1995, soil chemistry. Turner and Virtue (2006) and Holland-Clift 2004). The smothering Sorensen and Jusaitis 1995, Quarmby 2005, observed that dead rhizomes and tubers increases humidity around trees and pro- Willis et al. 2003, Taylor 2003, Downey were still present eight years after bridal vides optimal conditions for development 2006). Bridal creeper is likely to have ad- creeper had been killed. The decomposi- of diseases such as collar rot and Septoria verse impacts on biodiversity in other tion rate of bridal creeper’s below-ground fruit spot. It is estimated that at least 20% States where it is invasive, but these have biomass has not been investigated. In a of growers who manage a total of more not yet been recorded (Downey 2006). In preliminary experiment, an allelopathic than 6500 ha of citrus orchards in districts addition to the species listed in Downey response was obtained when bridal creep- bordering the Murray River from Mildura (2006), the orchid species Pterostylis bry- er roots were completely mashed up and to Cobram, are affected by bridal creeper ophila Jones in South Australia is also con- added to a hydroponic system, but these (Kwong and Holland-Clift 2004). Grow- sidered under threat due to habitat loss conditions were highly artifi cial and re- er’s perceptions of the negative impacts and bridal creeper invasion (Quarmby sults cannot be extrapolated to the fi eld of bridal creeper on their orchard ranged 2005). Furthermore, a recent assessment (A.J. Willis unpublished results). Howev- from increased labour and fi nancial costs, made in south-eastern New South Wales, er, it is noteworthy that root tissue of cul- damage to citrus trees, reduced fruit pro- which involved an extensive consultation tivated asparagus incorporated into soil duction, impediment to irrigation and process, suggests that up to 52 additional is reported to have allelopathic effects on fruit harvesting, and unsightliness. The plant species are potentially threatened by lettuce, tomato and asparagus seedlings cost of control is estimated to be as high bridal creeper invasion (Downey 2006). (Shafer and Garrison 1986). as $2000 ha-1 y-1 (Kwong and Holland-Clift The impact of bridal creeper invasion At Mt. Billy Conservation Park, a rem- 2004). In very severe infestations, where on threatened plant species can be dramat- nant eucalypt woodland in South Aus- >80% of the orchard is covered by bridal ic. For example, the density of the sand- tralia, greater accumulation of litter was creeper, growers believed that it is often hill greenhood orchid, Pterostylis arenicola observed in habitats invaded by bridal cheaper in the long run to remove trees M.A.Clem. & J.Stewart, at Tailem Bend, creeper compared to uninvaded native ar- and replant. South Australia was four times higher eas (Holt 2005, Stephens 2005). This differ- National expenditure on bridal creeper in plots where bridal creeper was absent ence was probably due to litter becoming control in natural ecosystems is impossible compared to infested plots (Sorensen and trapped in the climbing stems and foliage. to estimate, as many councils, community Jusaitis 1995). In south-eastern New South Increased litter accumulation can change groups and government agencies contrib- Wales, two of the largest and most ecologi- decomposition, nutrient and soil dynam- ute resources towards management of this cally signifi cant populations of the small ics, which in turn may affect establishment species and control cost per weed species shrub Pimelea spicata R.Br are in danger and growth of native species. Despite the is not necessarily recorded. The cost of re- of extinction primarily due to invasion by signifi cant decrease in plant diversity and search and implementation of the biologi- bridal creeper (Groves and Willis 1999, changes in habitat due to bridal creeper cal control program for bridal creeper was Willis et al. 2003). invasion, Stephens (2005) found a very estimated in 2002 to have reached at least Bridal creeper has also been identi- abundant and diverse arthropod and $6 million (Morin et al. 2006). fi ed as a threat to four endangered eco- wasp community in both native and brid- Bridal creeper is not known to be an logical communities in New South Wales al creeper invaded habitats. These study alternative host for diseases of economic (Downey 2006). Indeed it is believed to sites, however, were not completely colo- or environmentally important plants. Its have the potential to eliminate most un- nized by bridal creeper and consequently fruits are unpalatable to humans (Ray- derstorey species of invaded habitats the weed may not yet have caused a habi- mond 1999), but not reported as toxic. (Humphries et al. 1991). Downey (2006) tat change signifi cant enough to detect also identifi ed 11 additional ecological fl ow-on negative effects on the arthropod Benefi cial communities in New South Wales, includ- community (Stephens 2005). Bridal creeper has few redeeming quali- ing six already listed under the New South Bridal creeper’s possible detrimental ef- ties. It was widely grown as an ornamental Wales Threatened Species Conservation Act fects on native vertebrate fauna of natural plant from the mid 1880s to the early twen- 1995, that are potentially threatened by ecosystems have not been documented. tieth century, but its popularity declined bridal creeper. During the fruiting stage, established in- substantially afterwards. Today it is still Bridal creeper has also been shown to festations can harbour large numbers of sometimes seen growing in home gardens eliminate or restrict growth of a wide range exotic birds such as common blackbird (J. Hosking personal communication), al- of other native plant species. Turner and and starling (Raymond 1999, Stansbury though it is not sold anymore in nurseries Virtue (2006) demonstrated an increase in 2001), which are considered pests in Aus- and it is legislated against in all States. biomass and plant number of the native tralia (Bomford and Hart 2002). Although Bridal creeper’s fruits form part of the shrubs Rhagodia parabolica R.Br. and Rha- most frugivorous birds appear to prefer diet of a small number of native birds godia candolleana Moq., the groundcover fruits of native species over bridal creep- (Stansbury 2001), whose normal host Enchylaena tomentosa R.Br. and perennial er fruits, it is possible that availability of plants may occur in low density due to grasses in plots where bridal creeper was large quantities of the latter has a negative habitat degradation or destruction. Its foli- controlled over an eight year period. Simi- impact on the regeneration of native spe- age is non-toxic to mammals. It is reported larly, the number and abundance of native cies (Raymond 1999). to form a signifi cant part of the diet of tam- plant species was negatively correlated to Bridal creeper is not a weed of pastures mar wallabies (Macropus eugenii Desm.) on bridal creeper cover at sites in south-west- as it cannot withstand constant grazing Garden Island in Western Australia (Bell et ern Western Australia (P.J. Turner person- (Siderov and Ainsworth 2004). It is also al. 1987). Sheep and cattle have also been al communication) and South Australia not an invader in broadacre cropping situ- observed grazing on the foliage (Siderov (Stephens 2005). ations probably because of the limited seed and Ainsworth 2004), but there is no infor- Regeneration of native communities input in such environments, preference for mation on its nutritional value. after bridal creeper removal may be slow shaded habitats and the vulnerability of A higher number of soil/litter- where few propagules of native plants are seedlings to cultivation. Bridal creeper is, associated pollinators of the endangered left in the soil. Below-ground biomass may however, a troublesome weed of citrus orchid, P. bryophila, was recorded at bridal Plant Protection Quarterly Vol.21(2) 2006 55 creeper-invaded study sites in South Aus- Bridal creeper is a Class 1 plant in Herbicides tralia (Stephens et al. 2003, Stevens 2005). Queensland under the Land Protection (Pest Several herbicide trials have been con- It appeared that bridal creeper provided a and Stock Route Management) Act 2002, be- ducted in the last 20 years (Cooke and favourable habitat for these specifi c pol- cause it is not commonly present in the Robertson 1990, Pritchard 1991, 1995, 1996, linators and did not affect their movement State and has the potential to cause adverse 2002, McQuinn 1994, Dixon 1996), identi- or pollination effi ciency within the invad- economic, environmental or social impact fying glyphosate, metsulfuron methyl and ed habitat. (Anon. 2005i). It must be eradicated if it some related sulfonylureas as the most ef- In South Africa, bridal creeper is be- establishes in the State. It is declared a pro- fective non-selective systemic herbicides lieved to have therapeutical attributes. hibited plant (no sale or movement) in the against bridal creeper (Table 1). In a series A lotion prepared from its root biomass Australian Capital Territory under the Pest of pot trials, Pritchard (1991) found that is used to bathe sore eyes (Guillarmod Plants and Animals Act 2005 (Anon. 2006c). bridal creeper foliage was more sensitive 1971), although no detail on effi cacy and It is listed as a noxious weed to be eradi- to sulphonylurea herbicides than glypho- extent of use is provided in the litera- cated or not to be introduced to the North- sate, but that glyphosate efficacy was ture. Tubers have been seen being sold in ern Territory (class A and C weed) under improved with addition of the adjuvant the Cape Town market as a remedy for the Northern Territory Weed Management Pulse® (0.5% v/v). Increasing rates of these stomach problems, but again no detail is Act 2001 (Anon. 2006d). Bridal creeper has herbicides did not improve control in most known of the recommended method of recently been declared a restricted weed situations and especially when repeat ap- preparation or the dose rate (P.B. Edwards in Victoria under the Catchment and Land plications were made over consecutive personal communication). The chemical Protection Act 1994, banning its sale and years (Cooke and Robertson 1990, Pritch- composition of bridal creeper’s below and movement across the State (Anon. 2005j). ard 2002). Triclopyr/picloram, bromacil above-ground parts has not been invest- and a paraquat-amitrole-MSMA mixture igated. Weed management have also been reported to give good ini- As for other environmental weeds, lack of tial control in the fi eld, but less control was Legislation economic returns from control expendi- achieved with 2,2-DPA, 2,4-D/picloram, Asparagus asparagoides is prohibited for im- ture often precludes deployment of con- amitrole, triclopyr, tribenuron methyl and port to Australia under Commonwealth trol actions against bridal creeper at a local fl umetsulam, and no control was observed plant quarantine legislation (Quarantine scale. Use of traditional control methods with fl uroxypyr and MCPA/difl ufenican Act 1908) (Anon. 2005c). It is not permit- may also not be appropriate or feasible in (Cooke and Robertson 1990, personal ted entry into Western Australia under the some areas because of size and inacces- communications cited in Pritchard 1991, Plant Diseases Act 1974 and is declared a sibility of infestations, risk of off-target Yates 1997, Pritchard 2002). category P1 plant that cannot be sold, dis- damage when herbicides are used in con- Until recently, trials were generally tributed or moved within the State, under servation areas and labour-intensiveness carried out for a single year and did not the Agriculture and Related Resources Protec- of hand weeding because of the thick mat attempt to measure herbicide impact on tion Act 1976 (Anon. 2005d,e). Importation of rhizomes and tubers. These diffi culties bridal creeper below-ground biomass. into Tasmania is also restricted under the and the extensive Australian distribution However, Pritchard (2002) demonstrated Weed Management Act 1999 and the Plant of bridal creeper justify the substantial in- that a single application of glyphosate (180 Quarantine Act 1997 (Anon. 2005f). In that vestment that has been made to develop g 100 L-1) and Pulse® (0.5% v/v) reduced State, restrictions or measures for bridal effective classical biological control op- the living root system by 78% compared creeper are specifi ed in the weed manage- tions. to untreated plots. Between 90–99% of ment plan that is required under the Weed Prior to initiation of a control program, the root biomass was killed following a Management Act 1999 (Anon. 2005f). It it is useful to map the distribution and repeat application one or two years after must not be sold or distributed throughout densities of bridal creeper infestations in the initial application. In contrast, a single the State. Under the legislation, inspectors the targeted area or region (Carr 1996). application of metsulfuron methyl (1.5 g may require that land owners implement Such a map provides the basis for priori- 100 L-1) only reduced living root biomass measures to reduce the number of bridal tizing areas for control. Carr (1996) recom- by 32% and a repeat application after one creeper plants or eradicate or restrict the mends initially focusing control efforts on year of 1.5, 3 and 6 g 100 L-1 decreased weed from particular areas. low density infestations occurring in bush- the root system by 41–92%, 29–53% and Under the new South Australian Natu- land with the highest conservation value 74%, respectively (all rates with additive ral Resources Management Act 2004, bridal before tackling more seriously infested BS1000® (0.25% v/v)). Glyphosate (360 g creeper must be controlled and its trade areas. Targeting control towards climbing a.i. L-1; rate 20 mL L-1) combined with met- and movement are illegal throughout the plants that produce the most fruits within sulfuron methyl (Brushoff® 600 g a.i. kg-1; State (Anon. 2005g). In New South Wales the core infestation has been suggested to rate 0.1 g L-1) and a surfactant (LI700®; 5 bridal creeper is legislated under the Nox- limit spread (ARMCANZ et al. 2001). mL L-1) is currently used in the eradication ious Weeds Act 1993, and must not be sold, Since most bridal creeper seeds are campaign against bridal creeper on Lord propagated or knowingly distributed and bird-dispersed within 300–500 m of the Howe Island at readily accessible sites (Le the weed must be prevented from spread- source (Stansbury 2001, Siderov et al. Cussan 2006). ing to an adjoining property (W4c) in eight 2006), it is recommended to control outly- To achieve best results with chemical local council areas in the Sydney region ing plants or patches within a buffer zone control of bridal creeper, a spray program (Anon. 2005h). On Lord Howe Island it of 500 m around the edge of a main infes- of at least three years is recommended is listed under category W3 and must be tation (ARMCANZ et al. 2001). Siderov et (Robertson 1983, Cooke and Robertson prevented from spreading and its num- al. (2006) emphasized the importance of 1990, McQuinn 1994, Dixon 1996, Pritch- bers and distribution reduced. In the new controlling bridal creeper along paths and ard 1996, 2002). Repeat applications of Act implemented on 1 March 2006, bridal tracks within remnant vegetation because herbicides are essential to kill plants that creeper is banned from sale and move- they act as a conduit for invasion. Alterna- are missed in the fi rst year, any new re- ment (Class 5) for the whole of the State tively, reducing the number, or modifying growth and seedlings, and to have a sig- (D. Gannaway personal communication). the spatial arrangement, of tracks within nifi cant impact on below-ground biomass. Land holders are also required to control a reserve can be considered to limit seed A longer control program with herbicide the plant (Class 4) in local council areas dispersal within the area. applied every 2–3 years, rather than an- identifi ed in the Act. nually, may be a more effi cient use of time 56 Plant Protection Quarterly Vol.21(2) 2006 Table 1. Herbicides that provided signifi cant control (>90% when quantifi ed) of bridal creeper in different fi eld trials. Herbicide Formulation Rate Surfactant/Penetrant Comments Reference Glyphosate Glyphosate 360™ 360 g L-1 n/a Dixon 1996 (360 g a.i. L-1) 1 L ha-1, 1/100 360 g L-1, 1/50 n/a Graham and Mitchell 1996 360 g L-1, 1/70 n/a France 1996 180 g 100 L-1 With Pulse® (0.5% v/v) Higher rate and lower spray Pritchard 1996, 2001 spray vol.: 1000 L ha-1 volume did not improve results. More effect on root system than other herbicides. Metsulfuron methyl Ally® 2.5-5 g ha-1 in With BS 1000® May be more cost-effective Dixon 1996 Brushoff® 500–1000 L water (Alcohol alkoxylate) than glyphosate in areas (600 g a.i. kg-1) or Pulse® (modifi ed where the roots of non-target polydimethylsiloxane) plants are protected by a (0.2% v/v) with Ally good layer of litter. Brushoff 1.5 g 100 L-1 With BS 1000® (0.25% v/v) Higher rates and lower spray Pritchard 1996, 2001 spray vol.: 1000 L ha-1 or Pulse® (0.5% v/v) volume did not improve results. Thifensulfuron Harmony M® 15/1.5 g 100 L-1 With BS 1000® (0.25% v/v) Higher rate and lower Pritchard 1996, 2001 methyl/ (682/68.2 g a.i. kg-1) spray vol.: 1000 L ha-1 spray volume did not metsulfuron methyl improve results. Offered no improvement in root control over metsulfuron methyl. Chlorsulfuron Glean® 1.9 g 100 L-1 With BS 1000® (0.25% v/v) Offered no improvement Pritchard 1996, 2001 (750 g a.i. kg-1) spray vol.: 1000 L ha-1 in root control over metsulfuron methyl. TANK-MIXES Glyphosate and Glyphosate 360™ 90 g plus 1.5 g 100 L-1 With Pulse® (0.5% v/v) Lower spray volume did not Pritchard 1996 metsulfuron methyl and Brushoff® spray vol.: 1000 L ha-1 improve results These herbicides may not be approved for use on bridal creeper in all States/Territories and permits are required for off-label uses. Check with State or Territory weed management agency or local noxious weed authority for up-to-date information on registered herbicides and best application methods and dosage. and resources without compromising out- generally used to apply herbicides to brid- creeper was climbing. Several herbaceous comes (McQuinn 1994, Pritchard 1996, al creeper that is closely associated with species also survived after applications of 2002, ARMCANZ et al. 2001). Indeed, native vegetation and diffi cult to access metsulfuron methyl in Western Australian regrowth should be more abundant and with vehicles (Graham and Mitchell 1996). trials (Dixon 1996). Furthermore, some na- vigorous two years after initial applica- They are also ideal for spot-spraying iso- tive plants even survived after plots were tion, enabling higher uptake of herbicide lated plants, regrowth or new recruits that sprayed with glyphosate, probably be- and potentially more impact on the root emerge in years following initial herbicide cause they were partly protected from the systems. treatments or after a fi re (France 1996). spray by a dense bridal creeper canopy or Best control outcomes have been Techniques such as physically pulling off were at a growth stage that is more toler- achieved when metsulfuron methyl or bridal creeper foliage from surrounding ant to the herbicide. It may be more appro- glyphosate was applied onto actively native vegetation before spraying or wip- priate to use metsulfuron methyl instead growing bridal creeper in the early to ing instead of spraying have been sug- of glyphosate, even if it provides a lower late-fl owering stage (August-September) gested to minimize non-target effects from control of bridal creeper below-ground bi- (Dixon 1996, Pritchard 2002). Neverthe- applications of non-selective herbicides omass, in areas where it is not possible to less, good control was also obtained with (Carr 1996, Graham and Mitchell 1996, avoid spraying desirable ground species applications made at the fl ower bud to ARMCANZ et al. 2001). However, the in- (Yates 1997, Pritchard 2002). Nonetheless, green berry stage (Pritchard 2002). Spray- effi ciency and extra labour cost to imple- glyphosate can generally be sprayed onto ing before fl owering is advocated by some ment such techniques make them suitable bridal creeper climbing trunks of trees authors to allow better translocation to the only for areas of high conservation value without causing any non-target damage rhizome (Cooke and Robertson 1990) and (Graham and Mitchell 1996). (Graham and Mitchell 1996). Concerns to prevent off-target damage to indigenous The main challenge in using herbicides have been raised about possible adverse species that emerge in spring (Carr 1996). to control bridal creeper in natural ecosys- effects on native vegetation of long-term Effectiveness of glyphosate in killing tems is to avoid damage to non-target spe- use of metsulfuron methyl, which can bridal creeper is greatly enhanced when cies. In Pritchard’s (1996, 2002) fi eld trials, remain active in alkaline soils for many all foliage of an infestation is sprayed sedges and grasses present in treated plots months (ARMCANZ et al. 2001). It is thus (Dixon 1996, Graham and Mitchell 1996). were killed by glyphosate applications but recommended to avoid application of this Graham and Mitchell (1996) recommend not by metsulfuron methyl or other sul- herbicide near the root zone of trees (Dix- use of a marker dye in the herbicide mix fonylurea herbicides tested. In contrast, on 1996). and delineating grids using fl agging tape minimal or no damage was observed after Chemical control of bridal creeper in to ensure that no plants are missed or applications of any of the herbicides to fo- citrus orchards is not perceived by grow- only partly sprayed. Portable sprayers are liage of trees and shrubs on which bridal ers as being very effective, probably Plant Protection Quarterly Vol.21(2) 2006 57 because it cannot be used on bridal creeper underground rhizomes and tubers is in the burnt areas before regeneration of growing amongst citrus foliage due to the the main treatment method in the bridal sensitive native species (Carr 1996, France high risk of damage to citrus trees (Kwong creeper eradication campaign on Lord 1996, Graham and Mitchell 1996). Access and Holland-Clift 2004). For high value Howe Island (Le Cussan 2006). to infested areas for spraying is also con- citrus crops, trees can be ‘skirted’, where- siderably improved after a fi re because by lower limbs are pruned to enable under Slashing and grazing—Mechanical re- most woody shrubs have usually been tree access for spot-spraying with glypho- moval of bridal creeper above-ground bio- destroyed (Yates 1997). Willis et al. (2003) sate (ARMCANZ et al. 2001). mass is effective at reducing seed produc- suggested use of fi re in autumn, after most tion if performed before fruiting, but many annual shoots have emerged, and post-fi re Other treatments repeated slashings are necessary to exhaust application of herbicide on regrowth to Prevention—Education campaigns to in- stored underground reserves (Robertson maximize depletion of below-ground re- crease community awareness of problems 1983). Repeated mechanical defoliation of serves. Application of glyphosate follow- caused by bridal creeper are imperative bridal creeper in glasshouse experiments ing a prescribed burn has been found to be to keep un-infested areas free of the weed has been shown to reduce stored below- the most effective initial control treatment (ARMCANZ et al. 2001). Sale and dis- ground reserves (Raymond 1999, L. Morin against bridal creeper at two study sites in tribution of plants are prohibited across and A.J. Willis unpublished results). How- South Australia (Yates 1997). Although not Australia. Gardeners are encouraged to ever, plants totally defoliated every fi ve yet investigated in the fi eld, this strategy prevent further spread by avoiding plant- weeks for eight months still produced a could signifi cantly reduce bridal creeper ing or dumping of garden refuse contain- few new tubers and survived (Raymond infestations while stimulating germina- ing bridal creeper from their properties 1999). Slashing is sometimes carried out tion of native species such as P. spicata (ARMCANZ et al. 2001). Soil hygiene is in citrus orchards around trees infested by (Willis et al. 2003). There is no contra-indi- also important to prevent spread of vi- bridal creeper (Kwong and Holland-Clift cation to use biological control for bridal able rhizomes by earthmoving equipment 2004). Pruning of lower branches of trees creeper post-fi re. Indeed, biological con- (Cooke and Robertson 1990, Graham and is often required prior to slashing. trol agents are most likely to recolonize Mitchell 1996, Davies 2000). Bridal creeper is palatable to mam- sites if they are present in unburnt nearby mals. Tammar wallabies (Bell et al. 1987) areas (Morin et al. 2006). Hand weeding—Bridal creeper is eas- and quokka (Setonix brachyurus Quoy and ily recognized and isolated young plants Gaimard) (Carr 1996) are reported to eat Revegetation—Natural ecosystems may with an underdeveloped root system can bridal creeper foliage. On Garden Island take many years to recover after an area is be pulled-out by hand when growing in in Western Australia, Bell et al. (1987) ob- cleared of bridal creeper (Turner and Vir- loose soil (Carr 1996). Manual removal of served a reduction of more than 97% in tue 2006). Presence of bare ground after mature bridal creeper and its root system, bridal creeper cover in plots exposed to bridal creeper is controlled increases the a method often referred to as ‘grubbing’, is grazing by tammar wallabies (>100 indi- likelihood of re-invasion by bridal creeper only appropriate for small isolated infesta- viduals km-2) for two years compared to or other weeds. Land managers can ei- tions, particularly in areas of high conser- plots located within an enclosure. Slow ther rely on natural recovery processes or vation value (Robertson 1983, Cooke and invasion of bridal creeper from infested actively revegetate controlled sites with Robertson 1990, Carr 1996). It is also used tracks to the centre of a reserve on Phillip indigenous species. Pike (1996) observed by growers in citrus orchards who are Island, Victoria, may be due to feeding of natural regeneration of native plants, such reluctant to use herbicides (Kwong and swamp wallabies (Wallabia bicolor Desm.) as Hardenbergia comptoniana (Andrews) Holland-Clift 2004). Follow-up weeding within remnant vegetation (Siderov and Benth. and Acacia pulchella R.Br. in an area is often required because bridal creeper Ainsworth 2004). of the Boomerang Gorge where bridal regenerates from small pieces of living Livestock grazing is potentially an ef- creeper had been hand weeded. rhizome left behind in the soil (Pike 1996). fective control method for bridal creeper. Removed root mats should be disposed of Siderov and Ainsworth (2004) found no Natural enemies by deep burial (>2 m deep) or dried and bridal creeper occurring next to fenced ar- During 1996 and 1997, prior to the re- burnt (Graham and Mitchell 1996, Pike eas of pasture where it was abundant and lease of exotic biological control agents, 1996). supported the implementation of control- 34 bridal creeper infested sites in Western Hand weeding is time and labour in- led grazing in areas fenced off for revege- Australia (each of approximately 100 m2) tensive and consequently not suitable for tation to keep bridal creeper in check. Such were surveyed for presence of natural en- severe and extensive bridal creeper infes- an approach may be more effi cient than emies (including a sample of about 100 tations (Robertson 1983). Removal of well- regularly searching the area to remove leaves and fruits subsequently examined developed mats of rhizomes and tubers can newly-established seedlings (Siderov and in the laboratory) (K.L. Batchelor and J.K. create excessive disturbance that hinders Ainsworth 2004, Siderov et al. 2006), but Scott unpublished results). Very little leaf natural bush regeneration and facilitates may not be appropriate when the area is damage and no fruit or seed damage were weed invasion (Carr 1996). Carr (1996) revegetated with native species palatable observed. Raymond (1995, 1999) found emphasized that hand weeding should be to livestock. Cooke and Robertson (1990) only two phytophagous species feeding done only at the rate of natural regenera- reported use of sheep to reduce bridal on bridal creeper near one of her study tion to avoid over-clearing the area. creeper density prior to chemical control sites in Victoria: the weevil Phlyctinus callo- A minor bridal creeper hand weeding on Kangaroo Island, South Australia. sus Boheman and larvae of an unidentifi ed initiative was undertaken in the mid- Lepidoptera. Light brown apple moth (Ep- 1990s in the Boomerang Gorge, Yanchep Fire—Bridal creeper is not killed by bush- iphyas postvittana Walker) larvae were also National Park in an attempt to limit fur- fi res unless the fi re is very intense (Carr seen eating ripe fruit pulp but not dam- ther spread of the weed (Pike 1996). The 1996, Yates 1997). It is usually one of the aging seeds. Only one diseased specimen project was eventually abandoned be- fi rst species to emerge after a late summer of bridal creeper infected by a Phoma sp. cause of diffi culties in obtaining funding or early autumn wild fi re (France 1996, pathogen has been lodged in an Austral- to support volunteers’ activities and bet- Graham and Mitchell 1996). Land manag- ian mycological herbarium before biologi- ter control prospects with other methods ers are encouraged to take advantage of cal control was implemented (R. Shivas such as herbicide applications and biologi- a wild fi re and implement chemical con- personal communication). Localized snail cal control. Hand digging with removal of trol of emerging seedlings and regrowth (Helix aspersa Müller) damage has been 58 Plant Protection Quarterly Vol.21(2) 2006 observed on bridal creeper foliage in the 1999, Batchelor and Woodburn 2001). tactics such as fi re may be necessary in fi eld (L. Morin personal observations). Based on these results, the leafhopper, rust some areas to tackle bridal creeper more Field surveys were initiated in South fungus and leaf beetle were approved for effectively (Willis et al. 2003). No research Africa in the late 1980s to identify natural release in 1999, 2000 and 2002, respectively has yet been carried out to devise a practi- enemies of bridal creeper and determine (Morin et al. 2006). cal and effective strategy to integrate the distribution, seasonal occurrence and im- The rust fungus strain released in Aus- different control methods available. pact of those with potential for biological tralia, a single-uredinium isolate made control (Kleinjan and Scott 1990, Scott and from infected material collected at Jou- Acknowledgments Kleinjan 1991, Edwards 1995). Surveys bertsdal Farm near Swellendam in the We gratefully acknowledge CSIRO, the encompassed the range of bridal creeper winter-rainfall area of the Western Cape, Cooperative Research Centre for Weed in South Africa, but mostly concentrated only infects bridal creeper (Morin 1999). Management Systems that operated from on regions of the Western Cape Province The leafhopper can only complete normal 1995 to 2002 and the current Coopera- that are climatically similar to Australian development on bridal creeper, although tive Research Centre for Australian Weed infested areas. it can cause minor feeding damage on a Management for their continued support Natural enemies associated with bridal few other species within and outside the of studies relating to biological control of creeper in South Africa are discussed in Asparagus genus (Scott et al. 1999). Crop bridal creeper. The South Australian Ani- Kleinjan and Edwards (2006). Following asparagus and the only native Asparagus mal and Plant Control Commission and extensive excavations, bridal creeper tu- species in Australia, Asparagus racemosus the Australian and New Zealand Environ- bers and rhizomes were found to be rela- Willd., are not suitable hosts for the leaf ment and Conservation Council (previ- tively free of damage from natural enemies beetle (Batchelor and Woodburn 2001). ously CONCOM) were instrumental in (Scott and Kleinjan 1991, Edwards 1995). The biology, release, establishment and funding initial research on bridal creeper The weevil Brachycerus sp. (Curculionidae) initial impact of biological control agents biological control. Recently the National found on bridal creeper foliage may also released in Australia are discussed in de- Heritage Trust and the Department of feed on tubers in the larval stage but it has tail in Morin et al. (2006). Environment and Heritage have ensured still not been observed in the fi eld or labo- Although the rust fungus has already continuation of the program. We also ac- ratory (Kleinjan and Edwards 2006). Only demonstrated its reliability and superior- knowledge the hundreds of community two natural enemies, the moth Zalaca snel- ity in reducing density of bridal creeper members who have contributed their time leni (Wallengren) (Noctuidae) and larvae populations across most of its range in in the fi ght against this weed. We thank of the wasp Eurytoma sp. (Eurytomidae), Australia, it is likely to be complemented John Hosking (NSW Department of Pri- were observed feeding on fruits and seeds by the two invertebrate agents at some mary Industries), Richard Groves (CSIRO of bridal creeper (Edwards 1995, Kleinjan sites. For example, Turner et al. (2004) Plant Industry), John Virtue (Animal & and Edwards 2006). Eurytoma sp. was im- showed that a combined attack by the Plant Control Group Department of Water ported into an Australian quarantine fa- leafhopper and rust fungus resulted in an Land & Biodiversity Conservation Gov- cility for host range testing in 1999, but additive reduction to bridal creeper rela- ernment of South Australia) and Dane rearing diffi culties and host requirements tive growth rate. The leaf beetle, if it estab- Panetta (Queensland Department of Natu- when bridal creeper is not in seed remain lishes widely, should be active early in the ral Resources and Mines) for comments on to be resolved. Larvae of four moth spe- growing season before populations of the an earlier draft of the manuscript. cies belonging to the Noctuidae (Agrotis other agents build up (Morin et al. 2006). It sp., Euplexia augens Felder & Rogenhoven, may also have a main role to play in inland References Lycophotia oliveata (Hampson), Cucullia ter- areas where conditions are not optimal for Adair, R.J. and Groves, R.H. (1998). ‘Im- rensis Felder & Rogenhoven) were collect- the rust fungus. pact of environmental weeds on biodi- ed on bridal creeper foliage and reared in The biological control program against versity: A review and development of a the laboratory, but their specifi city has not bridal creeper has involved community methodology’. (Environment Australia, been investigated (Edwards 1995, Klein- groups, land managers and schools for re- Canberra). jan and Edwards 2006). An unidentifi ed distributing the leafhopper and rust fun- Agriculture and Resource Management cecidomyiid fl y (Cecidomyiidae) observed gus to new sites to enhance natural spread Council of Australia and New Zealand, producing galls on growing tips of bridal across Australia (Batchelor and Woodburn Australia and New Zealand Environ- creeper and other African Asparagus spp. 2002, Woodburn et al. 2002, Kwong 2002, ment and Conservation Council and and a foliage-feeding fl ea beetle, Hespera Batchelor et al. 2004). Some community Forestry Ministers. (2001). ‘Weeds of sp. (Chrysomelidae), were not investigat- groups have redistributed the rust fungus National Signifi cance: Bridal creeper ed further (Kleinjan and Edwards 2006). using a method referred to as the ‘spore (Asparagus asparagoides) strategic plan’. 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