94 Plant Protection Quarterly Vol.9(3) 1994 (AD); the Queensland Herbarium, Bris- The introduction and spread of capeweed, Arctotheca bane (BRI); the Australian National Her- barium, Canberra (CANB); the Tasma- calendula (L.) Levyns (Asteraceae) in Australia nian Herbarium, Hobart (HO); the Na- tional Herbarium, Melbourne (MEL); the Helen Wood, School of Science and Technology, Charles Sturt University, National Herbarium, Sydney (NSW) and Wagga Wagga, NSW 2678, Australia. the Western Australian Herbarium, Perth (PERTH). All specimens collected in Aus- tralia prior to 1985 were included; a full Summary listing of specimens is available from the Specimens of capeweed (Arctotheca ultimately recognized as weeds in the author on request. Of the 332 specimens calendula (L.) Levyns) from major Aus- new environment (Groves 1986b). This examined a small number were discov- tralian herbaria were examined in a information could be useful in a number ered to be misclassified (identified by Mr. study of the history of introduction and of respects; for example, by highlighting M. Gray as Arctotis sp. and Cymbonotus spread of capeweed in Australia. the methods which particular species use sp.) and some specimens in different her- Capeweed was introduced to Australia for long-distance dispersal and allowing baria were clearly duplicates, reducing from South Africa, probably on repeated quarantine procedures to be improved in the effective number of specimens. On the occasions, as a contaminant of stock fod- countries where the species have not yet other hand some sheets contained more der and packing straw. The earliest established. Studies of successful inva- than one plant (or parts of more than one record of its presence dates from 1834 sions may also indicate where control plant), on occasions collected at different (King George Sound, WA). measures could have been improved and times from a variety of locations, and Sites with early records of capeweed assist in preventing a repetition of past where possible these were considered as (Albany, Fremantle, Adelaide) have cli- mistakes when new species arrive. separate specimens in this analysis. For mates closely resembling that of Species for which information of this each specimen, the date of collection and Capetown, while capeweed took several type is known, such as subterranean clo- collector’s comments (if any) were re- decades to establish in climatically less ver (Trifolium subterranean) and Echium corded if available: flowering status and similar regions (e.g. Sydney). The rapid species (Piggin 1977, Gladstones and leaf form were also recorded for a sepa- spread of capeweed into the interior of Collins 1983), have complex introduction rate study (Wood 1993). The collection lo- the continent was aided by human trans- histories involving broad native ranges, cations of specimens were identified port; this trend commenced during the repeated introductions (both deliberate wherever possible and plotted in cells of expansion of agriculture in the 1800s and accidental), the likelihood of second- 1° latitude by 1° longitude on a map. and has continued unabated since then. ary introductions and genetically variable While capeweed is a weed of consider- parent populations, and in the case of Results able economic impact, it is regarded as a Echium a degree of taxonomic confusion useful component of pastures in low- and some differences of opinion about the History of introduction rainfall areas. The evidence suggests pattern of spread within Australia (Piggin The earliest dated specimen in this study that capeweed’s range in Australia is 1977, Kloot 1982). These complicating fac- (MEL105452) was collected in South Aus- still expanding. tors make it difficult to draw any conclu- tralia (Gawler Plain north of Adelaide) in sions about the relative importance of dif- 1848. However, capeweed arrived in Introduction ferent introduction routes or mechanisms Western Australia at least fourteen years Despite the best efforts of a competent of spread, or to make comparisons be- earlier, having been recorded from King quarantine service (Groves 1986b), exotic tween the species. George Sound (probably the vicinity of plant species continue to enter Australia This paper describes the history of in- Albany) in 1834 (Endlicher 1837) and Fre- undetected and to establish and natural- troduction and the pattern of spread of mantle in 1839 (Lehmann 1844); there is ize at an estimated rate of at least four capeweed (Arctotheca calendula) in Aus- also anecdotal evidence that the 1848 species per year (Specht 1981). Only a tralia. This species has a limited native specimen from Gawler postdates the ar- fraction of these plants are likely to create range being confined almost exclusively rival of capeweed in South Australia by weed problems (Groves 1986a), but meth- to the Cape Province of South Africa about ten years (Agricultural Bureau of ods for predicting which of the spectrum (Scott and Way 1990). Capeweed is a com- South Australia Report: Mount Gambier of invaders is a potential weed remain mon but relatively minor component of 1899). Introduction to Victoria and Tas- elusive. Climatic parameters may be use- the flora in its native range (Harvey and mania probably occurred before 1860 ful for predicting which weeds in other Sonder 1894, Adamson and Salter 1950), even though no specimens were collected countries would present similar problems but is a widespread and successful colo- from these States until 1886 and 1929 re- in Australia (Medd and Smith 1978, niser in southern Australia and is respon- spectively, since Hooker (1860) reported Panetta and Mitchell 1991), and investiga- sible for significant economic losses in that capeweed was abundant at Perth tions of factors such as geographical ori- both cropping and pastoral industries (Tasmania) and Hayter (1865) described gin (Forcella and Wood 1984, Fox 1987, (Shovelton 1982, Scott and Delfosse 1992). capeweed as common around Melbourne Kloot 1987), pattern and speed of spread The restricted native range and the likeli- in 1861. Herbarium records thus provide (Auld 1987, Forcella 1985) and various hood that capeweed was introduced di- an incomplete picture of the pattern of in- biological traits (Baker 1965, Newsome rectly from South Africa to Australia sug- troduction and spread of capeweed in and Noble 1986, Panetta 1987, Groves gest that the introduction history of Australia, especially in the early years of 1992) have attempted to identify an ‘inva- capeweed should be less fraught with European settlement. Despite the value of sive’ syndrome. However, it has not yet complications than the examples men- herbarium specimens in providing tangi- proved possible to predict the weed po- tioned above. ble evidence of the presence and spread tential of species which are not known to of introduced plants (Forcella and Harvey be weedy in other parts of the world. Methods 1983), such records must be supple- One aspect of plant invasions which Capeweed specimens from the following mented with information from other has received little attention is the history herbaria were examined: the State Her- sources when investigating patterns of in- of introduction of plant species which are barium of South Australia, Adelaide troduction and spread (Mack 1991). Plant Protection Quarterly Vol.9(3) 1994 95

a. 1875 b. 1895

c. 1945 d. 1985 Figure 1. Distribution of capeweed in Australia. Dots represent collection locations of herbarium specimens, stars represent locations based on other sources.

Capeweed was collected in New South Pattern of spread Within a matter of years after introduc- Wales (Port Jackson) in 1865 (MEL- Capeweed was introduced initially along tion several colonies developed at consid- 105466). This is probably an accurate in- the southern coast and as a consequence erable distances from the nearest estab- dication of the introduction date, since its subsequent spread has followed three lished coastal population (Figure 1a, b): there is no suggestion in the literature of general trends (Figure 1). The first of these specimens were collected from the its presence earlier than this date (Maiden was an inland spread from the coast away Murray River (NSW-Victoria border) in 1916b) and an annotation by the collector from the sites of introduction (Turner 1868, near Roma (Queensland) in 1871, indicates that it was a newly arrived spe- 1891). There was also a west-to-east Lake Bonney (SA) in 1874 and Eucla (WA) cies. Capeweed was reported as spread- spread from South Australia into Victoria and Omeo (Victoria) in 1882. This ing in the neighbourhood of Sydney in (Adcock 1915, Maiden 1916c) and a south- saltatory form of spread coincided with 1867 (Woolls 1867). Plants were collected to-north spread from Victoria to southern the rapid expansion of the pastoral indus- in Rockhampton in 1866, but it would ap- New South Wales and from New South try into the interior of the continent after pear that permanent populations have Wales into southern Queensland (Bailey about 1850, in the wake of explorers such failed to establish there; Bailey (1880) re- 1883, Agricultural Bureau of NSW Re- as Leichhardt, Sturt and Mitchell (Taylor ported noticing the species in Queensland port: Temora 1915, Maiden 1916c). West- 1990). about 1870, and by 1880 it was well estab- ern Australian populations have been The two decades between 1875 and lished along the Brisbane River (Bailey largely isolated from those in the eastern 1895 saw a substantial increase in collec- and Tenison-Woods 1880). States, but the same trends are evident. tions of capeweed (Figure 1b). This may 96 Plant Protection Quarterly Vol.9(3) 1994 be a result of the efforts of Baron von 200 30 Mueller and his collectors or may be a re- 180 flection of a sudden increase in capeweed 25 abundance or distribution. In either case, 160 by the turn of the century capeweed was 140 ■ ■ well established throughout most of Vic- ■ ■ 20 120 ■ toria, in the southern parts of Western and ■ ■ South Australia and in western and 100 ■ 123 15 123 1234 ■ coastal regions of New South Wales. 12■3 1234 ■ 80 1234 123 123■ 41234 1234 123 12341234 1234 123 12341234 10 Rainfall (mm) 12341234 123 12341234 Present distribution 60 Temperature °C 12341234 123 123412341234 As evidenced by herbarium collections, 12341234 123 123412341234 40 12341234 123 1234123412341234 capeweed is distributed widely through 12341234 123 1234123412341234 5 12341234 123 1234123412341234 20 12341234123412341234 123 1234123412341234 123 123 southern Australia (Figure 1c, d). In West- 12341234123412341234 123 1234123412341234 123 123

12341234123412341234 123 1234123412341234 123 123 ern Australia it is found throughout the 0 0 sheep-wheat areas of the south-west and Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec extends northwards along the coast as far a. Capetown, South Africa as Shark Bay. Although initially confined 200 30 to Adelaide and environs in South Aus- tralia, capeweed was first collected from 180 25 the Eyre Peninsula in the early 1960s and 160 has subsequently spread northwards into the interior of the State. Apart from the 140 ■ ■ 20 north-east corner, capeweed is found 120 ■ 1234 ■ ■ 1234 throughout Victoria. The previously dis- 123412341234 ■

123 123412341234 100 ■ 15 junct distribution in New South Wales 123 123412341234 ■ 123 123412341234 ■ ■ now covers most of the State and extends 80 123 1234123■ 4123■ 41234 123 1234123412341234 123 into south-eastern Queensland. It has 1234 123 1234123412341234 123 10

Rainfall (mm) 1234 123 1234123412341234 123 60 Temperature °C been recorded from Alice Springs in the 1234 123 1234123412341234 123 123 1234 123 1234123412341234 123 123 Northern Territory and Atherton in far 40 1234 123 1234123412341234 123 123 1234 123 1234123412341234 123 123 5 north Queensland. In Tasmania cape- 1234123412341234 123 1234123412341234 123 123 1234 20 1234123412341234 123 1234123412341234 123 123 1234 weed has not been collected from the 1234123412341234 123 1234123412341234 123 123 1234 1234123412341234 123 1234123412341234 123 123 1234 south-west but is present in more settled 0 1234 123 0 areas. Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec Although capeweed now generally oc- b. Albany, Western Australia cupies regions in Australia with similar 200 30 climatic patterns to those of its native 180 range in South Africa (Michael 1981), 25 160 specimens have been collected from loca- ■ ■ ■ tions which could be considered marginal 140 ■ ■ 20 at best. Early collections from Rock- 120 hampton in 1865 (AD 97943575, MEL ■ ■ 105455, MEL 105456, MEL 105472, MEL 100 ■ ■ 15 ■ 105477, MEL 105479, NSW 1377) and 1868 80 123■ 1234 123 123■ 41234 (MEL 105467) and from Atherton in 1947 123 12341234 10 1234 123 123412341234

Rainfall (mm) 60 Temperature °C (BRI 309502) have not been repeated sub- 1234 123 123 123412341234

1234 123 123 1234123412341234 sequently. Capeweed plants collected 40 12341234 123 123 1234123412341234 12341234 123 123 1234123412341234 123 5 near Alice Springs in 1962 (AD 96243306, 12341234 123 123 1234123412341234 123 1234 20 1234 12341234 123 123 1234123412341234 123 1234 AD 96349054, CANB 126906, NSW 1234 12341234 123 123 1234123412341234 123 1234 1234123412341234 123 123 1234123412341234 123 1234 NT9323, NSW NT9299) are clearly 0 1234123412341234 123 123 123412341234 1234 0 adventive (Chippendale 1972); of the two Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec recorded locations, one is adjacent to cat- c. Perth, Western Australia tle trucking yards and the other is a mown 200 30 123 (and presumably watered) lawn. Simi- 123 180 123 1234 larly all but one of the six South Austral- 123 1234 123 1234 25 ian specimens collected north of 30°S (AD 160 ■ ■ 123 1234 ■ 123 1234 96317102, AD 96519153, AD 97933330, 123 1234 ■ 140 123 1234 ■ 123 1234 ■ 20 AD 97934311, AD 97934346, AD 123 12341234

120 1234 123 12341234 ■ 97934369) were located at sources of wa- 123■ 4 123 12341234 1234 123 12341234 ■ ter (bore, spring, creek) or at stockyards 100 1234 12■3 12341234 15 ■ 1234 123 123■ 41234 or railway sidings. It is probable that col- 1234 123 12341234

80 1234 123 123412341234 lections from tropical Queensland and the 1234 123 123412341234

1234 123 123412341234 10

Rainfall (mm)

Temperature °C arid interior represent incidental and 60 1234 123 1234123412341234

12341234 123 1234123412341234 short-lived introductions; comparisons of 12341234 123 1234123412341234

40 12341234 123 1234123412341234 the respective climates with that of 12341234 123 1234123412341234 123 5

123412341234 123 1234123412341234 123 capeweed’s native range (Figure 2) sug- 20 1234123412341234 123 1234123412341234 123 123

12341234123412341234 123 1234123412341234 123 123 gest that the species is unlikely to survive 0 12341234123412341234 123 1234123412341234 123 123 0 in either environment in the long term. Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec d. Adelaide, South Australia Plant Protection Quarterly Vol.9(3) 1994 97 200 30 There is a possibility that capeweed was

180 1234 introduced from populations established 1234 1234 25 elsewhere, rather than direct from South 160 1234 1234 Africa. Since the evidence indicates that ■ 1234■ 1234 123 140 1234123■ 4 123 ■ capeweed did not become naturalized in 12341234 123 ■ 20 123412341234 123 ■ Europe until this century (Caballo 1940, 120 123412341234 123 ■ 123412341234 123 123 Tutin et al. 1976), secondary introductions 123412341234 123■ 123 ■ 123 100 123412341234 123 123 1234 123 15 from Europe could only have been as gar- 123412341234 123 12■3 ■ 1234 123 123 1234123412341234 123 123 ■ 1234 123 123 den specimens; therefore the balance of 80 1234123412341234 123 123 1234 1234 123 123 1234123412341234 123 123 1234 1234 123 123

Temperature °C probabilities favours the overwhelming 1234123412341234 123 123 12341234 1234 123 123 10

Rainfall (mm) 60 1234123412341234 123 123 1234123412341234 123 123 bulk of capeweed seeds introduced into 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 Australia having arrived as primary in- 40 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 5 troductions from South Africa. 1234123412341234 123 123 1234123412341234 123 123 20 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123

1234123412341234 123 123 1234123412341234 123 123 0 0 Routes of introduction Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec Anderson (1934) listed five possible intro- e. Sydney, New South Wales duction routes for weeds into Australia: 200 30 deliberate introductions, seed impurities, stock fodder, contamination in packing 180 ■ ■ ■ ■ ■ around goods and the ballast of ships.

1234 25

160 12341234 ■ ■ The relative importance of each route var- 12341234

12341234 ies depending on the particular life his- 140 12341234 ■ ■ 12341234 20 tory characteristics of the species in ques- 120 12341234 12341234 ■ ■ tion. In the case of capeweed all five 123412341234 ■ 100 123412341234 123 15 routes are possible, although some are 123412341234 123

123412341234 123 considerably more likely than others. 80 123412341234 123 123412341234 123 The most probable routes of introduc-

123412341234 123 123 Temperature °C

123412341234 123 123 10 Rainfall (mm) 60 tion are as seed contaminants in stock 123412341234 123 123 123412341234 123 1234 123 123 fodder and in packing material. The like- 40 1234123412341234 123 123 1234 123 123 1234123412341234 123 123 1234 1234 123 123 5 lihood is high that capeweed seeds were 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 present in hay loaded in Capetown to 20 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 feed livestock en route to Australia 0 1234123412341234 123 123 123412341234 123 123 0 Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec (Tench 1789). Fragile goods such as crock- f. Rockhampton, Queensland ery, glassware and wine were transported 200 30 in crates packed with straw, and this is ■ also a probable source of capeweed seeds ■ ■ 180 (Roach 1900, Maiden 1916c, Aplin and ■ 25 160 ■ Parsons 1988). ■ Other routes of introduction were less 140 20 ■ important. Capeweed was unlikely to 120 ■ have been brought into the country in bal- last initially since ships bound for Aus- 100 ■ 15 ■ tralia were fully laden with cargo; how- ■ 80 ■ ever after the 1850s when Australia be- 10 Temperature °C Rainfall (mm) 60 came a net exporter of agricultural pro-

1234 duce (Taylor 1990) seed in ballast may 40 123412341234 123 123412341234 123 5 have contributed to the subsequent accel- 123412341234 123 123 20 1234123412341234 123 1234 123 123 eration of spread. It is possible that 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 1234123412341234 123 123 capeweed was deliberately introduced for 1234123412341234 123 123 123412341234 123 123 0 0 ornamental purposes, having been in gar- Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec den cultivation in England since 1752 g. Alice Springs, Nothern Territory (Loudon 1880); Woolls (1867) suggested Figure 2. Climatic data for selected capeweed collection sites. Bars repre- that capeweed was introduced into the sent mean monthly rainfall, lines represent mean monthly temperature. Botanical Gardens at Melbourne, presum- Data for Capetown is from Takahashi (1986) and for Australian sites from ably as an ornamental species. This was Australian Bureau of Meteorology. probably of minor importance since plants growing along roadsides in Aus- Discussion and took on board cargo which provided tralia were viewed as novelties rather a vehicle for seed transport. For example, than as familiar garden escapes (Turner Sources of capeweed Captain Phillip obtained provisions as 1891). There is a suggestion that cape- During the early European colonization well as a variety of plants, seeds, livestock weed may have been introduced as a po- of Australia there were abundant oppor- and fodder in Capetown when the First tential fodder species (Anon 1880, cited in tunities for introductions from South Af- Fleet put in there in 1787 (Clark 1962), and Kloot and Burry 1982/83) and introduc- rica. Capeweed is a very common plant seven of the 21 ships entering Port tion as an impurity in seed for agricul- of roadsides and waste ground in its na- Jackson in the first nine months of 1801 tural crops is also possible. tive habitat (Adamson and Salter 1950, arrived from the Cape of Good Hope Harvey and Sonder 1894) and virtually (Aplin and Parsons 1988) while a further Establishment and climate every ship bound for Australia from Eng- six from London probably also stopped at A military garrison was founded at land made a victualling stop at Capetown Capetown on the way to Australia. Albany in 1826 and settlers arrived in 98 Plant Protection Quarterly Vol.9(3) 1994 1831 when the garrison was disbanded 120 (Jose 1930, Clark 1973), so capeweed was successfully introduced into King George 100 Sound within eight years of European oc- cupation and three years of civilian settle- ment. This surprisingly rapid establish- 80 ment probably reflects the similarity be- tween the climatic patterns of Capetown 60 and Albany (Figure 2a, b) and the next two sites where capeweed was recorded (Fremantle, near Perth, and Gawler, near 40 Adelaide) also have climates resembling that of its native range (Figure 2c, d). Cumulative number of cells 20 By contrast, colonization of Sydney dates from 1788 and there were ample op- portunities for capeweed to be trans- 0 ported there, yet the earliest specimen 1835 1845 1855 1865 1875 1885 1895 1905 1915 1925 1935 1945 1955 1965 1975 from the Sydney region (MEL 105466) Decade commencing dates from 1865, some 77 years later. The climate of Sydney (Figure 2e) has an ab- Figure 3. Spread of capeweed over time. Cells are 1° latitude by 1° longi- sence of the marked summer drought tude characteristic of Capetown and a higher “dirty chaff”, and contaminated fodder in cereal crops are recognized (Agricul- overall rainfall, and this difference in cli- was the seed source for new infestations tural Bureau of South Australia Report: mate may have hindered capeweed’s es- most often mentioned by collectors in this Kanmantoo 1898, Agricultural Bureau of tablishment on the eastern seaboard de- study. Other effective long-distance dis- South Australia Report: Boothby 1901, spite repeated opportunities for introduc- persal routes included horses, sheep’s Carne 1926) and tainting of milk is also tion. Commenting on the absence of wool and packing material (Agricultural reported (Agricultural Bureau of South capeweed from coastal New South Wales, Bureau of NSW Report: Temora 1915, Australia Report: Mount Gambier 1899, Tenison-Woods (1887) speculated that Maiden 1916c). Pierce 1970), but it is also perceived as capeweed may require dry windy The pattern of spread after introduction “often more beneficial than injurious” weather during summer to ensure disper- to a new area was more reminiscent of an (Agricultural Bureau of South Australia sal and encourage autumn germination; advancing front, suggesting that different Report: Murray Bridge 1898, Carne 1926). summer rainfall was suggested as a limit- dispersal methods were more important Apiarists value capeweed as a pollen ing factor to this establishment. for local spread. These include wind dis- source for honeybees (Clemson 1985) and Morphological variation among cape- persal, contaminated lawn turf and the it even seems to have been deliberately weed herbarium specimens indicates that transplantation of roadside plants into sown to provide fodder for sheep and populations in New South Wales may be gardens (Turner 1891, Maiden 1916a, c). pigs (Agricultural Bureau of South Aus- genetically different from their counter- Roadsides and waste ground were impli- tralia Report: Mount Gambier 1899). parts in other States as a reflection of cated as important sources of infestation, The dichotomy of opinion regarding separate introductions (Wood 1993). Al- with capeweed reported as spreading capeweed’s value is related to land use ternatively, it is possible that the along roadsides (Ewart 1913, Maiden and climate. It is regarded as a weed in capeweed biotypes which eventually es- 1916c, Carne 1926). crops generally and in pastures in higher- tablished in Sydney were secondary in- rainfall regions, while in drier areas it is troductions derived from an Australian Is capeweed a weed? considered to make a valuable contribu- population in which ecotypic adaptation Although capeweed has been identified tion to pastures (Carne 1926, Scott and was occurring (Wood and Degabriele as a serious weed in both crop and pas- Way 1990). 1985); after 1850 Albany was the first ture situations (Shovelton 1982, Velthuis landfall for ships bound for the eastern and Amor 1982), an ambivalence towards Future prospects for control ports (Appleyard 1981), so it is possible its weed status is evident (Scott and Way The relatively short lag phase after intro- that some establishments after this date 1990). In New South Wales its weed po- duction and the high initial rate of spread involved capeweed derived from Albany tential was recognized as early as 1890 displayed by capeweed (Figure 3) are (or some other Australian source) rather (Turner 1890, 1892); by 1915 capeweed typical of weeds which have a wide final than directly from Capetown. Capeweed was rated seventh in a list of the 20 worst distribution (Forcella 1985). Migration was among a collection of 60 plant spe- weeds in the State and had been declared rates of many introduced plant species cies obtained from King George Sound noxious in 37 shires and municipalities are best described by a sigmoid curve during a three-hour stay (Anon 1890), in- (Maiden 1915). The number of declara- (Auld et al. 1982/83, Forcella 1985) and dicating its proximity to an effective tions had increased to 47 by 1934 the curve for capeweed continues to rise means of eastward migration. (Anderson 1934). Similarly, in Victoria steeply so it is probable that this species capeweed was first proclaimed in 1901 has not yet reached the limits of its distri- Methods of spread (Ewart and Tovey 1907) and within 15 bution in Australia; the apparent rapidity Rapid spread over long distances is typi- years was one of the commonest weeds in of its spread through the Eyre Peninsula cal of species in which seed dispersal is the valley of the Upper Murray (Ewart in the 1960s is a case in point. The poten- mediated by human activity (Panetta 1913). In a statewide survey of Victoria in tial distribution of capeweed in Australia 1987), and transportation of stock fodder 1978 capeweed was rated as the most im- could be predicted by homoclime analy- is a likely route (Thomas et al. 1984). portant weed of lucerne and pasture in sis (Panetta and Mitchell 1991) and Maiden (1916c) stated that the appearance Victoria (Shovelton 1982). uninfested areas thus identified should of capeweed in the New England district Opinions about capeweed appear to be be carefully monitored in future. The of New South Wales after the drought of more equivocal in South Australia and potential for biological control by 1902 was attributed to the importation of Western Australia. Its detrimental effects phytophagous insects is currently under Plant Protection Quarterly Vol.9(3) 1994 99 investigation (Scott and Way 1990, Scott South Wales 4 (ser 1), 26-36. Subterranean clover as a naturalized and Delfosse 1992). However, the history Bailey, F.M. (1883). ‘A synopsis of the plant in Australia. Journal of the Austral- of failed efforts to eradicate, or even con- Queensland flora; containing both the ian Institute of Agricultural Science 49, trol, introduced weeds (Medd 1987, phanerogamous and cryptogamous 191-202. Streibig 1992) leads to the conclusion that plants’, p. 267. (Government Printer, Groves, R.H. (1986a). Invasion of weeds capeweed will continue to spread in Aus- Brisbane). in Mediterranean ecosystems. In ‘Resil- tralia. Bailey, F.M. and Tenison-Woods, J.E. ience in Mediterranean climate ecosys- (1880). A census of the flora of Brisbane. tems’, eds. B. Dell, A.J.M. Hopkins and Acknowledgements Proceedings of the Linnaean Society of B.B. Lamont, pp. 129-45. (Dr. W. Junk, I thank the Director and staff of the Aus- New South Wales 4 (ser 1), 137-204. The Hague). tralian National Herbarium, Canberra, Baker, H.G. (1965). Characteristics and Groves, R.H. (1986b). Plant invasions of for arranging the loan of herbarium speci- modes of origin of weeds. In ‘The ge- Australia: an overview. In ‘Ecology of mens on my behalf, and in particular Max netics of colonizing species’, eds. H.G. biological invasions: an Australian per- Gray for expert advice and assistance. Baker and G.L. Stebbins, pp. 147-68. spective’, eds. R.H. Groves and J.J. (Academic Press, New York, USA). Burdon, pp. 137-49. (Australian Acad- References Caballo, A. (1940). ‘Flora analytica de emy of Science, Canberra). Adamson, R.S. and Salter, T.M. (1950). Espana’. (Sociedad Anonima Espanola Groves, R.H. (1992). Weed ecology, biol- (Eds.) ‘Flora of the Cape Peninsula’, de Traductores y Autores, Madrid, ogy and spread. Proceedings of 1st In- pp. 828-9. (Juta and Co, Capetown, Spain). ternational Weed Congress, 17-21 Feb- South Africa). Carne, W.M. (1926). Cape Weed. Journal ruary 1992, 1, 83-88. Adcock, G.H. (1915). Rutherglen Experi- of the Department of Agriculture of West- Harvey, W.H. and Sonder, O.W. (1894). ment Farm: Survey of the commoner ern Australia 3, 480-1. ‘Flora capensis: being a systematic de- weeds. Journal of Department of Agricul- Chippendale, G.M. (1972). Check list of scription of the plants of the Cape ture Victoria 12, 337-46. Northern Territory plants. Proceedings Colony, Caffraria and Port Natal’. Vol- Agricultural Bureau of New South Wales of the Linnaean Society of New South ume III, pp. 466-8. (L. Reeve and Co, Report: Temora. (1915). Agricultural Ga- Wales 96, 207-67. Brook). zette of New South Wales 26, 820. Clark, C.M.H. (1962). ‘A history of Aus- Hayter, H.H. (1865). Remarks on a Agricultural Bureau of South Australia tralia’. Volume I, p. 85. (Melbourne filamentous material, grown upon the Report: Boothby. (1901). Journal of Agri- University Press, Melbourne). plant “Cryptostemma Calendula- cultural Industry of South Australia 5, Clark, C.M.H. 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