140 Protection Quarterly Vol.20(4) 2005 transport organisms (Planty-Tabacchi et al. 1996, Stohlgren et al. 1998). Here we re- Invasion history and ecology of the environmental port on the history of the introduction and weed balloon , Cardiospermum grandiflorum colonization of one such invader, balloon vine (Cardiospermum grandiflorum Swartz; Swartz, in Australia ) in Australia. While recog- nized as an environmental weed for at Scott P. CarrollA,B, Michael MathiesonC and Jenella E. LoyeA,B least fifty years, it has received formal at- tention only recently (Batianoff and Butler A Department of Entomology, University of California, Davis, CA 95616, USA. 2002). Batianoff and Butler (2003) rank it B 13th in terms of prospective environmen- Department of Zoology and Entomology, University of Queensland, St. tal impact among 66 ‘priority’ environ- Lucia, Queensland 4072, Australia. mental weeds in south-east Queensland. C Environmental Protection Agency, Wildlife Ecology Unit, 80 Meiers Road, It is one of the 26 ‘Class 3’ species listed for Indooroopilly, Queensland 4068, Australia. Queensland (Land Protection Act 2003). These are declared pest the sale of which is illegal. We are particularly inter- ested in balloon vine because of its incur- Abstract from ornamental plantings in parks and sions into threatened rainforest habitats in The tropical American balloon vine, Car- gardens (Martin 2002). For nearly all Aus- eastern Australia, and because specialized diospermum grandiflorum, is declared tralian weeds there is a lack of historical insects that attack its in its Neotropi- as an environmental weed throughout information on dates of introduction and cal homeland have close relatives here Queensland and in many districts of rates of spread, deficiencies that will need that may play a role in limiting its rate of New South Wales. Using Australian her- to be overcome if our ability to predict spread (Carroll et al. in press). barium records we documented its intro- future negative impacts is to be refined As structural parasites, that grow duction and spread, mapped its invasion, (Groves et al. 2003). into the upper canopy may have dispro- described its habitats, and documented Naturalization is a key element of the portionate impacts on native tree-domi- its shift from domestic to natural envi- invasion process. If factors influencing the nated communities. Notably, several of the rons and from garden plant to recognized probability of naturalization are substan- most serious environmental weeds in rain- threat. Balloon vine was introduced into tially different in Australia than in the tem- forest areas of Queensland and New South Sydney, northeastern New South Wales perate Northern Hemisphere, case studies Wales are indeed vines (e.g., Madeira vine, and southeastern Queensland in the of invasions on this continent will be an Anredera cordifolia, cat’s claw creeper, Mac- 1920s and 30s from unknown sources. especially valuable component of control fadyena unguis-cati, and climbing aspara- Range expansion was relatively slow science here. Such documentation should gus fern, Protoasparagus africanus. Nine until the 1960s, and major increments include analyses of a taxon’s origin, resi- (35%) of the declared Class 3 pest plants have occurred in the past 15 years. It dence status, and invasion status (Pysek for Queensland (Land Protection Act 2003) was collected near Adelaide in the 1980s, et al. 2004). Data on invasion after intro- are vines. Large vines such as balloon vine and Perth in the 1990s. Naturalization duction can give insight into the process- may be regarded as ‘transformer’ species is evident in some of the earliest collec- es that underlie successful establishment because of their ability to markedly alter tions. Local eradication efforts occurred and potentially aid in control (Williamson ecosystem properties (Vivian-Smith and as early as 1935, and its weed status was 1996). A large proportion of invaders may Panetta 2002, Pysek et al. 2004). Thus, the recognized in some regions by the 1960s. be ‘sleeper weeds’ that begin to spread control of these plants is of particular con- It grows on a diversity of soil types and only decades or centuries after introduc- servation interest. Balloon vine is especial- is particularly abundant in riparian cor- tion (Groves 1986, Kowarik 1995), sug- ly recognized as a threat in southeastern ridors, where it may cover other vegeta- gesting that a variety of environmental Queensland, northern New South Wales, tion in uninterrupted stands kilometres and genetic factors may come into play and in the vicinity of Sydney (Batianoff in length. Much of its inland spread is in determining an invader’s subsequent and Butler 2002, Muyt 2001) and below. along watercourses. spread and abundance (Hobbs and Hum- Key words: alien, balloon vine, ecol- phries 1995, Crooks and Soulé 1999). Chief Native distribution ogy, exotic species, history, naturalized, among these may be natural or anthropo- Cardiospermum grandiflorum is widely dis- plant invasion, Sapindaceae, weed. genic habitat alteration (e.g., Hobbs 1991, tributed in its native Neotropical range, Hobbs and Atkins 1988) and adaptive occurring from southern Mexico to Brazil Introduction evolution (Lee 2002). These ecological and and the Caribbean (e.g., Cowan 1983, Bal- Environmental weeds are nonindigenous evolutionary aspects of invasion biology ick et al. 2000, Brako and Zarucchi 1993, plants that invade natural habitats. In may be examined by reconstructing the in- Moraes 1990). The type specimen is from Australia, the transformation of alien troduction and spread of an environmen- Jamaica. For 180 specimens in the Missouri plants to environmental weeds is recog- tal weed. For example, notes on the habitat Botanical Garden and New York Botanical nized as problem of great ecological and or ecology commonly accompany botani- Garden databases, the elevational range is economic significance (e.g., Humphries et cal specimens in herbaria. Such records 10–1800 m. In South America it occurs in al. 1991, Csurhes and Edwards 1998, Mar- may reveal the extent to which an invader the Caribbean lowlands, at low to mid el- tin 2002). Of approximately 28 000 plant is dependent on disturbed habitats, or the evations in moister areas of the equatorial species brought into Australia in the past chronology by which it began to colonize Andes, in southwestern Amazonia, and in 200 years, close to 10% have become envi- natural (versus domestic) environments. Paraguay and southern Brazil. In Belize, ronmental weeds (Martin 2002, Groves et Among native plant communities, ripar- Central America, it occurs commonly on al. 2003). This frequency greatly exceeds ian habitats, which typically harbour high river and stream banks at low elevations, the conventional ‘1% rule’ for naturaliza- native species diversity, may be particu- while in Costa Rica it also occurs on forest tion that stems from temperate zone stud- larly prone to invasions. This vulnerability edges at roadsides up to more than 1000 ies (e.g., Kowarik 1995). An estimated two- may stem from their physically dynamic m elevation (SPC personal observation). thirds of Australia’s weeds are escapes nature, high nutrient levels and ability to Populations in South and Oceania Plant Protection Quarterly Vol.20(4) 2005 141 are nonnative and of conservation concern (Olckers 2003, Meyers 2002, respectively). Seeds are dispersed by wind and water (Harden 2002). In the Neotropics, seeds of C. grandiflorum and its congeners are at- tacked by a variety of insects (Carroll and Loye 1987, Carroll 1988, Carroll and Loye in press). Little else is known of its biology and ecology.

Materials and methods We investigated the history of balloon vine’s incursion and spread within Aus- tralia by examining 122 specimens from the National Herbarium (17 specimens), Queensland Herbarium (48 specimens), National Herbarium of New South Wales (44 specimens), University of New Eng- land Beadle Herbarium (seven specimens), and the North Coast Regional Herbarium � � � � � � � � � � in Coffs Harbour, New South Wales (six specimens). Both cultivated and natural- ized plants were present. Where duplicate specimens were held among herbaria, only a single occurrence was tabulated. All specimen data from the Western Australian Herbarium were duplicated in the National Herbarium, and that from the CSIRO Atherton Her- barium was duplicated in the Queensland � Herbarium. No specimens were found in the herbaria of the Australian National � � � � � � � � � � � � � � � University, James Cook University, or the Northern Territory. We mapped the plant’s history of spread based on places and dates of collections. The goal of mapping was to give a clear sense of the time course of geographic � ���� expansion. Where data are few, as in the earliest collections, we mapped the points � ���� individually. We grouped later collections ����������� into four intervals of 15 years. The manner � � in which we depicted the spatial extent ����������� of each 15 year grouping was based on two assumptions. The first is that some of ����������� the outlying points are derived from geo- graphically intermediate populations that ����������� have not yet been discovered or reported. The second is that many of the known locales are but single sites in widespread Figure 1. Map of the invasion of eastern Australia by balloon vine, metapopulations. This approach may im- Cardiospermum grandiflorum, since 1923. See text for collections from South ply the existence of interstitial populations Australia and Western Australia. for which we lack collection data, but re- veals the time course of change in a manner that individually-mapped records cannot. A majority of the locales we used are avail- South Wales, in 1923. The species is absent a 1949 specimen taken 150 km north of able on the worldwide web at Australia’s from the early floristic treatments of New Sydney at Singleton (Hunter River Valley) Virtual Herbarium, and the complete data South Wales by Dixon (1906) and Maiden was from the fence of a hotel. Garden in- set is available from the first author. and Betche (1916). The second herbarium troduction is a likely source of the vine Lastly, at several points we make refer- record for is also from the Sydney region, in both southern (Beadle et al. 1962) and ence to general observations on the extent in Lane Cove in 1935. In that year an- northern New South Wales, and its con- of infestations made in field surveys con- other specimen was collected 600 km to tinued cultivation may in part underlie the ducted in 2002–2004. the north in Kyogle (Figure 1). Additional coalescence of the northern and southern New South Wales specimens were collect- distributions in the 1960s (Figure 1). Results ed adjacent to these foci in the subsequent Early floras of Queensland did not Introduction to Australia decades. In notes on a specimen collected reference the plant (e.g., Bailey 1909). The earliest herbarium specimen of C. at Bellvue Hill (near Sydney) in 1954, E.J. Hill (1875) lists the small congeners C. grandiflorum in Australia was taken from a McBarron remarked that the vine was ‘cul- halicacabum and C. h. var. microcarpum in river bank near Sydney in Camden, New tivated in many country towns’. Likewise, the holdings of the Queensland Botanic 142 Plant Protection Quarterly Vol.20(4) 2005 Gardens (in Brisbane) at that time, but Habitat, soil and phenology For collections in Queensland there are makes no mention of C. grandiflorum. The Of the 122 collections, 79 included infor- far fewer status notations. Details do not earliest Queensland records are both in the mation about the habitat of the specimen, appear until the 1990s, with notes such as southeast of that state: at Laidley near Gat- or strong clues thereto. Fifty-two of these ‘vigorous’, ‘abundant’, ‘rampant’, ‘smoth- ton in 1944, and at Coolum in 1945. While collections were reported from along wa- ering trees’, and ‘climbing over canopy 16- not far removed from northern NSW latitu- tercourses, and another six were almost 20 m’ appear with increasing frequency. It dinally, two factors suggest that they may certainly so, yielding a total of 70% ripar- was collected in Lamington National Park be separate from the Kyogle introduction. ian. Another two were from gullies, and (Kerry Valley) in 1991. We observed it at First, they are on the opposite side of the a third from the ‘margin of mangrove 1900 m elevation in Springbrook National Border Ranges, which should form a natu- swamp’, further indicating a predilection Park in 2004. ral barrier to water and wind dispersal of for this type of environment (77%). The re- The two collections from South Austral- the seeds. Second, they are both adjacent maining records consist of roadsides (6%), ia simply refer to the vine as ‘vigorous’. In to the greater Brisbane region, a likely area waste sites (5%), cultivated or suburban Western Australia, the vines along the Can- for horticultural plantings, with potential (5%), open forest (3%), and disturbed for- ning River were described as ‘rampant… for escape. ests (4%). 8–10 m over trees… locally abundant’. Subsequent expansion of the range was Soil types varied substantially among most pronounced in the 1960s, and from the collected specimens. Among the 11 Discussion the 1990s to the present (Figure 1). The first records were four alluvial, two gravely, Cardiospermum grandiflorum is a canopy- substantially inland records occurred in and one each skeletal, dark grey, basalt, climbing vine that is native in warm habi- the early 1960s in Queensland at Emer- rich loamy and mud sandy loam. This re- tats from southern Mexico to southern ald. In 1969, a specimen was collected in sult indicates that a diversity of soil types Brazil. It is not well studied in its native South Australia in Adelaide, and again in is suitable for the growth of balloon vine. region, but supports a number of eat- the same region a decade later. Few col- There appears to be a latitudinal in- ing insects (Carroll and Loye 1987, Car- lections took place beyond the established fluence on reproductive seasonality. In roll and Loye in press). Due to its curious range perimeters in the 1970s and 1980s, NSW, all records mentioning flowers or seedpods and attractive appearance, it is but since 1990 many collections have been fruit took place February–July. In contrast, planted as an ornamental in many warm made both inland and farther north in Queensland records indicate fruiting and areas of the world. In addition to its decla- eastern Australia (Figure 1). In addition, flowering in spring and summer in ad- ration as an environmental weed in many specimens were taken along the Canning dition to autumn and winter. The more areas of eastern Australia, it is a very seri- River in Perth, Western Australia, in 1995. equably warm climate of the north may ous environmental weed in South Africa On a state-by-state basis, the herbaria support a longer breeding period on an (Olckers 2003) and the Cook Islands (Mey- hold a total of 59 specimens from NSW, 58 annual basis. ers 2002). from Queensland, two from SA and one The geographic origin of the C. gran- from WA. (Two additional specimens from Evidence of pest status and control diflorum imported into Australia is un- ACT are from cultivated specimens at the The collection records indicate that bal- known. There appear to have been at least National Botanic Gardens, originally from loon vine was recognized as a weedy pest five Australian foci for its introduction. Queensland stock.) Major portions of east- species early on. For example, the 1935 Residence time on the continent is at mini- ern Australian habitats now either support specimen from a gully in Lane Cove was mum 80 years. We treated the three main balloon vine populations, or are close to from an individual that was ‘poisoned’. east coast introductions in some detail, such populations. E.J. McBarron’s collection notes from the and reported introductions at Adelaide greater Sydney region offer the most de- and Perth. While data are lacking, there History of naturalization tailed observations on the status of bal- is substantial indication that horticultural We searched for the starting point of loon vine. In addition to his notes about its plantings of this species were popular naturalization in collection notes accom- naturalization and cultivation in 1954, he historically, and as we know of no other panying the herbarium specimens. Feral noted it climbing over trees and ‘proving practical reasons for its introduction, we individuals appear early in the collection troublesome’ where cultivated by 1965. In suspect that balloon vine was originally a history. As noted above, the earliest collec- 1965, at Wooloomooloo, another collec- garden escape. tion (1923) is from a ‘river bank’. The next tor annotated a specimen with ‘An order The large size and distinctive appear- note is for the Lane Cove NSW specimen, for the destruction of this vine has been ance of C. grandiflorum suggest that its ab- taken from a ‘gully’ in 1935. A 1947 record served on the owner’. In a collection at sence from collections prior to 1923 is not from Longueville (near Sydney) is from Windsor in 1971 it was simply described a result of its having been overlooked. As an ‘open eucalypt forest’. In 1951 it was as a ‘weed’, and a 1977 specimen includes a contrast, its comparatively diminutive ‘in the bush and on roadside’ in nearby a newspaper article about the spread of and uncommon congener C. halicacabum Wahroonga, and McBarron’s 1954 note re- the weed in Fairfield (near Sydney). Speci- was noted in Bentham (1863), and was ports the species in that region as ‘over mens from the 1980s and 1990s include frequently collected from the 1800s on in scrub, also naturalized at Cheltenham’. notes of ‘Over trees to 9 m’ and ‘forming both Queensland and NSW, including in The first habitat characterization for vine towers’. areas from which C. grandiflorum was later northeastern NSW is from Ulmarra in In far northeastern NSW, the Rich- collected. As a result, we surmise that the 1957, where the plant was described as mond River near Kyogle currently sup- early herbarium collections give a good ‘wild on roadside’. Six of the seven speci- ports many linear kilometres of balloon chronology of the plant’s introduction. mens collected in this region in the 1960s vine arrayed along its floodplain (SPC From among the first collections, there were from river banks (see below for more personal observation). However, in 1964, is evidence that balloon vine began repro- results on habitat). E.F. Constable described the population ducing outside of cultivation early in its Habitat notes are not present for in that region as ‘localized on rainforest tenure on the continent. The great majority Queensland specimens before 1968. Eight edge…’ At the same site four years later, of herbarium specimens are from non-cul- of 12 specimens collected from that time A.G. Floyd described it as ‘commonly tivated plants, and many of them are from through the 1970s were from river banks, growing over trees and shrubs…’ By 1990 bush lands, especially riparian habitats. indicating a naturalized status by that G.J. White described it as ‘vigorous’ in the While the rate at which specimens are point in time. rainforest there. taken has increased in the 80 years span, Plant Protection Quarterly Vol.20(4) 2005 143 the proportion that is from noncultivated no balloon vine in cursory surveys of is attributed to a tropical cyclone (Meyers and wild habitats has been high through- that valley, and in similar surveys in 2004 2000), and Brisbane area naturalists com- out. Continued horticultural plantings found just four individuals, none of them ment on the great spread of balloon vine may have seeded new areas of escape all large. The impression, then, is that even in in that region following the major cyclone through this period. long-occupied regions, certain catchments and flooding of 1974 (D. Sands personal We lack information on the exact places (perhaps those with comparatively high communication). Furthermore, even if and dates at which C. grandiflorum was in- elevation margins such as the Tweed) are substantial evolution has not attended its troduced in various areas of Australia. As still in early stages of colonization. initial invasion of Australia, subsequent a result, it is difficult to determine whether As balloon vine colonizes westward evolution may improve its ability to colo- there was any substantial lag time between along watercourses, wind dispersed prop- nize other habitats. Such a process would introduction and naturalization. Data agules may become even more important be facilitated by its now massive popula- showing that lags of up to 150 years are in the colonization of new ‘oasis’ habitats tion size, as well as its continued ecologi- commonplace (e.g., Kowarik 1995) make capable of supporting the plant. The much cal release from directly adapted parasites it tempting to speculate that balloon vine smaller congener C. halicacabum occurs and competitors. Once established, eco- was introduced well before the 1920s. Such patchily in central and northern coastal logical transformers such as balloon vine a ‘pre-naturalization’ period would have Queensland, and more commonly in mon- may perennially inhibit recolonization by increased the number of available prop- soonal areas of NT and WA. Based on its natives. Accordingly, the invasion of Aus- agules and better-adapted genotypes, and native range, it is likely that C. grandiflo- tralia by this Neotropical exotic deserves provided time for response to any strong rum has both the adaptive and dispersive continued attention and control efforts. selection (sensu Hobbs and Humphries ability to ultimately occupy these habitats 1995, Crooks and Soulé 1999). as well. Acknowledgments On the other the other hand, studies By their capacity to enter the canopy, For sharing their knowledge of balloon showing both lags and low establishment invasive vines may be particularly detri- vine and Queensland environmental rates under natural conditions (i.e., the ‘1% mental to their host communities. Balloon weeds, we are particularly indebted to G. rule’, Kornas 1990, Kowarik 1995) are all vine has been recognized as troublesome Batianoff (Queensland Herbarium) and from temperate sites. Establishment rates for at least 60 years. Current infestations G. Vivian-Smith and D. Panetta (Alan in Australia are closer to 10% (Martin 2002, appear massive to the extent that floral Fletcher Research Station). For extensive Groves et al. 2003). High establishment fre- and faunal relations are probably deeply and generous logistic and editorial sup- quencies may indicate that the native flora altered, although this has not been meas- port, we thank M. Zalucki and H. Dingle is comparatively invasible or that there ex- ured. Listing as a Class 3 pest in Queens- (University of Queensland). L. Jessup, W. ists a large global pool of subtropical spe- land means that selling any plant material MacDonald, P. Robins, and H. Winters cies pre-adapted to prevailing abiotic and without permission is illegal. However, (Queensland Herbarium), M. Garcia (Roy- edaphic conditions here. If either is the such a measure is unlikely to directly im- al Botanic Gardens, Sydney), R. McKin- case, lag periods during which substantial pact its further colonization and establish- non (Brisbane Botanic Gardens), A. Floyd evolution takes place before naturaliza- ment. Local legislation is exercised in an (Coffs Harbour Herbarium), J. Hauser, tion can commence may not typify many increasing number of districts, and has the R. Whyte and members of The Hut En- of the invasions in Australia, and balloon greatest prospect for controlling balloon vironmental Centre (Brisbane) assisted in vine may actually have moved out of the vine, particularly in and near cities and the search for sites and records of balloon garden early on. The multiple foci of its towns. A Google internet search of ‘bal- vine. The Australian-American Fulbright invasion reinforce this interpretation of loon vine Australia’ reveals many local Commission and Carroll-Loye Biological its endogenous invasive power in eastern organizations actively working against Research funded this work. Australian habitats. it in New South Wales and Queensland. The seeds of Cardiospermum are dis- The Canning River population that was References persed by both water and wind (Harden the source of the 1995 Perth collection ap- Bailey, F.M. (1909). ‘Comprehensive cata- 2002). Mature fruits that remain attached to pears to have been eliminated by local logue of Queensland plants both indig- the plant dehisce and release seeds carried bush regeneration efforts as of late 2001 enous and naturalized’. (A.J. Cumming, on specialized wings. Fruits may also drop (SPC and JEL personal observations). We Brisbane). intact, and float readily on water (Vivian- are not aware of extensive naturalization Balick, M.J., Nee M.H. and Atha D.E. Smith and Panetta 2002). This two-tactic in SA at present. (2000). Checklist of the vascular plants dissemination strategy serves to permit The invasion of riparian and adjacent of Belize. Memoirs of the New York Bo- thorough colonization within and between habitats by balloon vine may be espe- tanical Garden 85, i-ix, 1-246. watersheds. Our findings indicate that cially threatening to declining rainforest Batianoff, G.N. and Butler, D.W. (2002). soil type is relatively immaterial. Presum- communities in eastern Australia (sensu Assessment of invasive naturalized ably, the colonization of new watersheds Randall 1986, Planty-Tabachi et al. 1996, plants in south-east Queensland. Plant requires propagules to cross topographic Martin 2002). We have observed numer- Protection Quarterly 17, 27-34. divides, an act best accomplished by air. ous areas in which the vine has spread Batianoff, G.N. and Butler, D.W. (2003). In recent field surveys (unpublished) we from creek banks into forest, particularly Impact assessment and analysis of six- noted interesting heterogeneity in the satu- in sites with some physical disturbance ty-six priority invasive weeds in south- ration of adjacent watersheds by balloon (SPC and JEL personal observations). In- east Queensland. Plant Protection Quar- vine. For example, the herbarium collec- vasions are often triggered by rare events terly 18, 11-17. tion data indicate that in northeastern New (Rejmánek et al. 2004). Landscape level Beadle, N.C.W., Evans, O.D. and Carolin, South Wales, the Richmond River catch- disturbances such as hurricanes may open R.C. (1962). ‘Handbook of the vascular ment has supported balloon vine for at least and deposit seeds in habitats from which plants of the Sydney district and the 50 years and probably longer. Currently this plant was previously excluded. In ad- Blue Mountains’. (Published by the au- many reaches of that catchment (including dition to specialized structures for wind thors. Armidale). tributaries) appear dramatically ‘smoth- dispersal, balloon vine fruits are capable Bentham, G. (1863). ‘Flora Australiensis’, ered’ by balloon vine. In contrast, there are of floating in water for months (Vivian- Volume 1. (Reeve and Co., London). no collections from the large Tweed River Smith and Panetta 2002). The invasion of Brako, L. and Zarucchi J.L. (1993). Cata- Valley just to the north. 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