Cereus uruguayanus (Cactaceae) and its naturalised occurrence in Queensland, Australia Paul I. Forster & Miriam Schmeider Summary Forster, P.I. & Schmeider, M. (2000). Cereus uruguayanus (Cactaceae) and its naturalised occurrence in Queensland, Australia. Austrobaileya 5(4):671-677. The first records of Cereus uruguayanus Ritt. ex Kiesl. as a naturalised weed in Australia are reported. Several populations occur in western Queensland on heavy clay soils in natural and disturbed woodland of brigalow (Acacia harpophylla F.Muell. ex Benth.) and belah (Casuarina cristata Miq.). Endozoochorial dispersal is thought to be responsible for the spread of this species in natural vegetation and eradication is recommended. It is estimated that at least 3240 individuals occur at one locality near Glenmorgan. Size class structure of this population is described which shows a preponderance of seedling juveniles and large mature plants. The stand is also notable for the high proportion of fasciated (10.9%)and monstrous (20.2%) individuals that occur. This represents the first numerical data on fasciation and monstrousity in a population, albeit naturalised, of Cactaceae. Keywords: Cereus uruguayanus, fasciation, monstrousity, naturalised weeds P.I. Forster & M. Schmeider, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,Toowong QLD 4066 Introduction Species of Cactaceae have been remarkably uruguayanus Ritt. ex Kiesl. encountered during successful in colonising the Australian a visit in February 1997 to the property “Myall continent with thirty species currently Park” near Glenmorgan. An additional considered as being naturalised (Forster 1996). naturalisation has also been found near Tara The majority of naturalised species are from and another reported from the gemfields at the genus Opuntia, although taxa from the Anakie (J.Higgins pers. comm. 1999). genera Acanthocereus, Epiphyllum, Harrisia (syn. Eriocereus), Hylocereus, Nyctocereus, Cereus uruguayanus is native to Pereskia and Selenicereus are also present Argentina, Brazil and Uruguay but has been (Telford 1984; Hosking etal. 1988; Forster 1996). widely cultivated around the world since the early 1800’s, usually under the name The majority of these naturalised species C.peruvianus Mill. Kiesling (1982) established are shrubby to arborescent, spiny succulents that the name C. peruvianus was misapplied with Opuntia tomentosa Salm-Dyck attaining and renamed the species as C. uruguayanus a height of 7 m in some situations. To date, Ritt. ex Kiesl. Hunt (1992) has referred without cacti with globular (eg. Echinopsis or justification, both the names C. peruvianus Mammillaria) or candelabra habits (eg. Cereus auct. and C. uruguayanus to the synonymy of and related genera) have been largely absent C. hildmannianus and Taylor (1998) has from the Australian naturalised cactus flora. recently recombined C. uruguayanus as a Naturalisations of species such as Echinopsis subspecies of C. hildmannianus. This latter multiplex (Pfeiff) Zucc. have been localised combination was made in a privately published adventives and easily contained (Mann 1970; journal series that specialises in automatic Hosking et al. 1988) and were excluded from tranfers of names with often no justification to recent listings of naturalised cacti (eg. Telford support them. In the case of Taylor’s new 1984; Forster 1996). In this paper we document combination there is no explanation offered and an extensive naturalisation of Cereus until such time as a comprehensive revision of the genus is provided it is more appropriate to follow the nomenclatural lead of Kiesling (1982), Accepted for publication 5 January 2000 a recognised authority on Argentinian cacti. 672 Austrobaileya 5(4): 671 - 677 (2000) In his classic work on the cultivation of Weber 1966; Hunter 1984; Sauleda & Sauleda cacti, Borg (1937) stated that the species [as 1984) and often formally named (eg. Borg 1937; C. peruvianus] was “Long known in Lindsay 1963; Backeberg 1976). We believe our cultivation”. Despite its ubiquity in cultivation, study to be the first that details the numerical there is little ecological or taxonomic occurrence of fasciation and monstrosity, albeit information available about this species. Britton in a naturalised population. & Rose (1920) in their monograph of Cactaceae Materials and Methods provide a brief account of both C. hildmannianus and C. peruvianus, stating Site Description & History. The study site is that the former occurs in Brazil and the latter in situated on the property “Myall Park” (27°12’S, south-eastern South America. Benson (1982) 149° 39’E), near Glenmorgan some 330 km west commented that the species (as C. peruvianus) of Brisbane. “Myall Park” is the site of a private was commonly naturalised on Kauai in Hawaii botanic garden “Myall Park Botanic Garden where it was sometimes a pest of pastures. Ltd.” that is primarily devoted to Western Most contemporary books on cacti omit Australian species and was established in the mention of the species (eg. Barthlott 1979; 1940’s by the redoubtable David Gordon Andersohn 1983) and accurately identified (McKenzie 1995). Up until the introduction of illustrations are scarce and generally incomplete the moth Cactoblastis cactorum (Berg) in 1933, lacking flowers and fruit (Taylor 1968; Hunter the area around Glenmorgan was severely 1988a; Glass & Foster 1989; Innes & Glass 1991; infested with 'prickly pear’ (Opuntia stricta Silva & Sazima 1995). At a locality in south¬ (Haw.) Haw.) and would appear to be suitable eastern Brazil, C. uruguayanus is stated to for the persistence of different sorts of cacti. occur on rocky outcrops in both forested and Several other species of cacti were believed to deforested areas (Silva & Sazima 1995, as have been cultivated at the “Myall Park” C. peruvianus). These authors found that this homestead by the late 1940’s (N.Lester, pers. cactus was predominantly pollinated by comm. 1997) and it is assumed that the hawkmoths and that seasonal flowering population of C. uruguayanus originates from coincided with an activity peak for these this time. C. uruguayanus was semi- insects. commercially available from nurseries by 1936 Fasciation and monstrosity of the stem (Swinbourne 1982) and by the late 1950’s was is common in naturalised Australian commonly cultivated inAustralia (Fuaux 1957; populations ofC. uruguayanus, hence we also Hayes 1958 [all as C. peruvianus]). report on its numerical occurrence at one locality. Fasciation in cacti occurs when the At “Myall Park” individuals of apical meristem divides in an abnormal manner C. uruguayanus (Voucher: Forster 20334 & forming unusual fan-shaped stems (Synder & Watson: BRI) are concentrated (27° 12' 16MS, Weber 1966; Boke & Ross 1978; Gibson & 149° 39’ 35"E) in c. 6 ha of disturbed, but Nobel 1986) and is thought to be due to several reasonably intact woodland dominated by factors, such as external stimuli, disease or Casuarina cristata Miq. (belah) and Acacia heredity (Synder & Weber 1966). Monstrosity harpophylla F.Muell. exBenth. (brigalow) on in cacti occurs where each shoot loses its heavy clay alluvium. Brigalow often occurs as vegetative point after producing a few areoles ‘clumps’ as a result of the formation of‘gilgai’, with new growth points produced in an irregular which are depressions in the soil that hold manner. Such fasciated and monstrose clones water after heavy rain (Johnson, 1980). The of cacti are often popular as ornamentals. To cacti are predominantly concentrated in these the best of our knowledge, there are no detailed clumps of brigalow. Scattered individuals occur studies of fasciation and monstrosity in natural outside of this area, nearly always in clumps of populations of cacti, although the occurrence natural vegetation, and the furthest individual of isolated individuals in the wild is observed was about 1 km away from the main occasionally reported as a curiosity (eg. Graham naturalisation (27°12'40"S, 149o40'14"E). A 1962; Lindsay 1962; Foster 1965; Synder & similar distribution of individuals was also Forster, Schmeider, Cereus uruguayanus in Queensland 673 . Fig. 1 Cereus uruguayanus. A. budding ‘normal’ stem. B. ‘monstrous’ stem. C. ‘cristate’ stem. D. fruit (whole). E. fruit cross-section. F. seedling. All from Forster 20334 & Watson (BRI). Del. W. Smith. 674 Austrobaileya 5(4): 671 - 677 (2000) observed in the population nearTara (Voucher: ‘NormaF individuals make up the bulk of the Forster PIF24959 & Booth, BRI), except that total population surveyed (68.9%), followed by the brigalow community is more disturbed those with cristate stems (20.2%) and being mainly regrowth. monstrous stems (10.9%) (Fig. 3). ‘Cristate’ individuals are more frequent in the smaller size Methodology: Ten quadrats of 10 x 5 m were classes and for plants over 2 m in height make placed deliberately within the main up only 9.3% of the population (Fig. 2). concentration of cacti. All individuals of cacti in a quadrat were scored for several features of Discussion size class and stem form. Natural History Four size (age) classes of plants were designated - There are few detailed studies available of size 0-50 cm (classified as juveniles) (Fig. IE). class structure in cacti and none for natural populations of Cereus uruguayanus. As 50-100 cm (classified as mature as they are indicated in the materials and methods, the capable of flowering and several showed cacti were noticeably concentrated in natural evidence of this). vegetation of brigalow clumps. In South Africa, 1 -2 m (generally unbranched if with normal stem Taylor & Walker (1984) found that the closely morphology). relatedC. jamacaru DC. [as C.peruvianusbut see Glen 1997 for correct nomenclature] could > 2m (generally branched and with a candelabra only establish on fine-textured soils with a high habit). density of shade trees. The requirement of “prey refugia” and “nurse” plants that create a Three classes of stem organisation were recognised - suitable microclimate for establishment of succulent plants is now well known ‘normal’ stems (Fig. 1 A) where the 5-7 ribs are (Steenbergh & Lowe 1969; Nobel 1988; not sinuately indented between areoles McAuliffe 1984). The clumped distribution of and are more or less straight. C. uruguayanus at “Myall Park” indicates that a similar process is occurring, but it is likely to ‘monstrous’ stems (Fig. IB) where 7 or more be mainly due to “nurse” plant availability ribs are present with marked indention rather than “prey refugia”. Most of the between areoles and the ribs are rarely seedlings observed were not hidden in dense straight. natural vegetation, and predation was ‘cristate’ stems (Fig. 1C) where it is not possible noticeably absent on individuals that were to accurately ascertain the rib number due otherwise readily accessible. Brigalow clumps to the form of apical cell division where may act as a “nurse” plant for C. uruguayanus many areoles are densely concentrated by providing microclimatic conditions suitable and the ribs are never straight. for seedling establishment and by acting as foci for seed dispersal. Spiral stems as illustrated by Hunter (1988a) were not observed nor were cristate flowers as This cactus is dependent on cross¬ described and illustrated by Muller (1988). pollination between different individuals for Results: Two hundred and sixty-seven fruit to be produced (Silva & Sazima 1995). The individuals of C. uruguayanus were recorded resultant fleshy fruit with numerous seeds (Fig. from the 10 quadrats with an average of twenty- ID) appears suited for endozoochorial dispersal seven individuals per plot. Based on this by birds (Bregman 1988). Most of the cacti that average it is estimated that the total population are serious pests in Australia are thought to could be in excess of 3240 individuals. have fruit (and hence seed) that are eaten and Collectively there was a preponderance of dispersed by birds and mammals (Hosking et immature seedlings and large (> 1 m) individuals al. 1988). If this is the case forC. uruguayanus, (Fig. 2). it would be worth observing birds that utilise Forster, Schmeider, Cereus uruguayanus in Queensland 675 Size classes Fig. 2. Total size class distribution for 267 individuals of C. uruguayanus in 10 quadrats at “Myall Park”. brigalow clumps for roosting to determine those although in instances where they were closely that feed on these fruit and whether they situated competition for resources would had disperse seed over any distance. to have been a factor. Once established, growth of seedlings of C. uruguayanus is rapid and As yet this naturalisation is relatively localised, maturity is reached within 3 or 4 years (pers. but isolated plants up to a kilometre away would obs. 1978-1997 on cultivated plants at Didcot). indicate successful endozoochorial dispersal Prior to 1996 the area near Glenmorgan had is occurring. The brigalow belt, although experienced over 5 years of periodic drought widespread in eastern Australia, is now and this may have been responsible for a lack endangered due to clearing for intensive of intermediate sized plants that would have agriculture and cattle grazing and has about established in that period. The only way to 2.2% of its original occurrence conserved in determine these sorts of trends would be to reserves (Young et al. 1999). Given the scale of establish permanent plots with tagged the naturalisation of C. jamacaru in South individuals; however, in the current situation it Africa by 1984 (c. 3000 ha), and the success of would be better if the population was other cacti as agricultural and environmental eradicated. weeds in Australia (Hoskingef al. 1988; Forster 1996), particularly in brigalow communities (McFadyen 1984), it is important that it be monstrous successfully controlled. cristate 11% The size class distribution of individuals at “Myall Park” is different to that found by Taylor & Walker (1984) for C. jamacaru as there is a greater preponderance of seedlings and large mature individuals in relation to intermediate sized plants. Such a ‘bell’ shaped distribution was implied by Taylor & Walker (1984) to indicate unstable populations where stand structure had not yet stabilised and competition between individuals was not restricting seedling establishment. At “Myall Fig. 3. Total percentage of ‘normal’, ‘cristate’ and ‘monstrous’ individuals of 267 C. uruguayanus Park”, seedlings were generally well scattered, plants in 10 quadrats at “Myall Park”. 676 Austrobaileya 5(4): 671 - 677 (2000) Fasciation naturalised population of C. uruguayanus is probably unusual for the relatively high Fasciation and monstrosity in C. uruguayanus percentage of such plants but may be a result was first documented by de Candolle in 1800 of the founding individual or individuals for cultivated material and various infraspecific carrying genes for this abnormality. Hunter taxa have been named to encompass these (1988b) stated that seedlings from fruit of the forms (Britton & Rose 1922, Borg 1937; Kiesling ‘monstrose’ form of C. uruguayanus will be 1982). These infraspecific taxa have been nearly 100% true to fonn. In the current example, mainly described under C. peruvianus and juveniles or small adults of both ‘cristate’ and none have been transferred to C. uruguayanus ‘monstrous’ individuals were more common to date and most discussion of the species still than large mature individuals over 2 m in height. persists under the misapplied name (eg. Hunter 1988a,b; Muller 1988, Machadoet al. 2000). Acknowledgements This current study demonstrates that fasciation Thanks are due to W. Smith for the drawings in and monstrosity may arise continuously within Fig. 1, to A.Wood for translation of Kiesling a population indicating a genetic tendency for (1982), to N.Lester and D.Gordon for snippets this to occur. It has been speculated that of Myall Park history, to C.C.Walker for a copy somatic cross-overs are a possible mechanism of Taylor (1998) and to G.N.Batianoff and for inducing this variation (Machado et al. P.D.Bostock for reviewing the manuscript. 2000). Some ‘normal’ individuals of References C. uruguayanus were also noted as having the occasional ‘monstrous’ or ‘cristate’ branch, Andersohn, G. (1983). Cacti and Succulents. Wakefield: hence there is no justification for recognition EP Publishing Limited. of such teratological forms as infraspecific taxa as undertaken by Britton & Rose (1922) or Backeberg, C. (1976). Cactus Lexicon. 3rd ed. Poole: Blandford Press Ltd. Backeberg (1976). Rather, if such forms have to be provided with a name, then selected clones Barthlott, W. (1979). Cacti. Cheltenham: Stanley should be designated as cultivars. Given the Thornes (Publishers) Ltd. confused history and doubtful typification of the infraspecific taxa for fasciated individuals Benson, L. (1982). The Cacti of the United States and Canada. Stanford: Stanford University Press. described under C. peruvianus (Kiesling 1982), it would be wise to arrive at a totally new set of Boke, N.H. & Ross, R.G. (1978). Fasciation and names for such forms of C. uruguayanus if so dichotomous branching in Echinocereus required. (Cactaceae). American Journal of Botany 65:522-530. Fasciated individuals of cacti occur rarely in Borg, J. (1937). Cacti. London: Macmillan & Co. nature (eg. Graham 1962; Lindsay 1962,1963; Ltd. Synder & Weber 1966) or cultivation, but seem to be very commonly recorded for Bregman, R. (1988). Forms of seed dispersal in Cactaceae. Acta Botanica Neerlandica C. uruguayanus^Kiesling 1982; Hunter 1988a; 37:395-402. Muller 1988; Glass & Foster 1989, Machado et al. 2000). In the case of the monstrous forms of Britton, N.L. & Rose, J.N. (1920). The Cactaceae. Lophocereus schottii (Englem.) Britt. & Rose Descriptions and Illustrations of Plants of the Cactus Family. The Carnegie Institute of described by Lindsay (1963), both were Washington, Publication 248, Vol. 2. thought to be clones that reproduced Washington. vegetatively. Both Graham (1962) and Lindsay (1962) stated or inferred that some populations Forster, PI. (1996). Naturalised succulents in the of cacti tended to have a greater tendency for Australian flora. Haseltonia 4:57-65. fasciation than others, but apart from Foster, R. (1965). A tortuous form of Machaerocereus mentioning some localities, did not document gummosus. Cactus & Succulent Journal (Los numerical occurrence in the wild. This Angeles) 37:158-159. Forster, Schmeider, Cereus uruguayanus in Queensland 677 Fuaux, L. (1957). Questions & Answers. Cactus & Mcauliffe, J.R. (1984). Prey refugia and the Succulent Journal of New South Wales 1(3):32. distributions of two Sonoran Desert cacti. Oecologia (Berlin) 65:82-85. Gibson, A.C. & Nobel, P.S. (1986). The Cactus Primer. Cambridge: Harvard University Press. Mcfadyen, R.E. (1984). Alien weeds. In A.Bailey (ed.): The Brigalow Belt of Australia, pp. 177-182. Glass, C. & Foster, R. (1989). Dear Abbey Garden Brisbane: The Royal Society of Queensland. Press. Cactus & Succulent Journal (Los Angeles) 61:39-40. Mckenzie, B. (1995). One Man’s Dream. David Gordon and Myall Park Botanic Garden. Glen, H.F. (1997). Cactaceae. In Smith, G.F. et al. Glenmorgan: B.McKenzie & Myall Park (eds.): List of Southern African Succulent Botanic Garden Ltd. Plants, pp. 49-50. Pretoria: Umdaus Press. Muller, H. (1988). Cristata-blute bei Cereus Graham, D. (1962). In quest of crests. Cactus & peruvianus (Linne) Miller. Kakteen und andere Succulent Journal (Los Angeles) 34:156-158. Sukkulenten 39:111. Hayes, R. (1958). Editor’s Medallion. Cactus & Nobel, P.S. (1988). Environmental Biology of Agaves Succulent Journal of New South Wales 1(7):91. and Cacti. Cambridge: Cambridge University Press. Hosking, J.R., Mcfadyen, R.E. & Murray, N.D. (1988). Distribution and biological control of cactus Sauleda, R.P. & Sauleda, D.K. (1984). A crested form species in eastern Australia. Plant Protection cf Echinocereus chloranthus (Engelm.) Quarterly 3:115-123. Rumpler var. chloranthus from Texas. Cactus & Succulent Journal (Los Angeles) 56:243. Hunt, D.R. (1992). Cites Cactaceae Checklist. Kew: Royal Botanic Gardens. Silva, W.R. & Sazima, M. (1995). Hawkmoth pollination in Cereus peruvianus, a columnar Hunter, M. (1984). Another look at fasciating cacti. cactus from south-eastern Brazil. Flora Cactus & Succulent Journal (Los Angeles) 190:339-343. 56:40-42. Swinbourne, R.F.G. (1982). The history of cacti and Hunter, M. (1988a). Spiralling cactus. Cactus & succulents in Australia, 1787-1981. Calandrinia Succulent Journal 60:87-88. 2: 36-41. Hunter, M. (1988b). Collecting crested and monstrose Synder, E.E. & Weber, D.J. (1966). Causative factors Euphorbias. Euphorbia Journal 5:20-25. of cristation in the Cactaceae. Cactus & Innes, C. & Glass, C. (1991). Cacti. New York: Succulent Journal (Los Angeles) 38:27-32. Portland House. Taylor, N.P (1998). Nomenclatural adjustments in various genera of Cactaceae (subfam. Johnson, R.W. (1980). Studies of a vegetation transect through brigalow (Acacia harpophylla) forest Cactoideae). Cactaceae Consensus Initiatives 6: 15-16. in central Queensland. Australian Journal of Ecology 5:287-307. Taylor, R. (1968). Cactus growers guidelines. Cactus Riesling, R. (1982). Problemas nomenclaturales en el & Succulent Journal (Los Angeles) 40:131. genero Cereus (Cactaceae). Darwiniana Taylor, S.E. & Walker, B.H. (1984). Autecology of 24:443-453. an invading population of the cactus Cereus Lindsay, G (1962). Giant crests. Cactus & Succulent peruvianus (Queen of the night) in the central Journal (Los Angeles) 34:172-174. Transvaal. South African Journal of Botany 3:387-396. Lindsay, G. (1963). The genus Lophocereus. Cactus & Succulent Journal (Los Angeles) 35:176-192. Telford, I.R.H. (1984). Cactaceae. In A S.George (ed.): Flora of Australia 4:62-80. Canberra: Australian Machado, M.F.P.S, Mangolin, C.A. & Collet, S.A.O. Government Publishing Service. (2000). Somatic crossing-over can induce isozyme variation in somaclones of Cereus Young, P.A.R., Wilson, B.A., Mccosker, J.C., Fensham, peruvianus Mill. (Cactaceae). Haseltonia 7:77- R.J., Morgan, G. & Taylor, P.M. (1999). 80. Brigalow Belt. In P.Sattler & R. Williams (eds.): The Conservation Status of Queensland’s Mann, J. (1970). Cacti naturalised in Australia and Bioregional Ecosystems. Brisbane: their control. Brisbane: Department of Lands. Environmental Protection Agency, Queensland Government.