128 Plant Protection Quarterly Vol.21(3) 2006 dioecious species examined had on aver- age twice the genome size of the South Af- Review of the current taxonomic status and rican hermaphroditic species and, based on internal transcribed spacers of nuclear authorship for Asparagus weeds in Australia ribosomal DNA, the species can be divided into two clusters, one of European species Kathryn L. Batchelor and John K. Scott, CSIRO Entomology, Private Bag 5, and the other of southern Africa species. PO Wembley, Western Australia 6913, Australia. Both Lee et al. (1997) and Stajner et al. Email: [email protected], [email protected] (2002) did not use outgroup taxa in their analyses. This issue and a larger sam- ple size (24 species) were addressed in a study by Fukuda et al. (2005) of the mo- Summary Systematic taxonomy of genera lecular phylogeny of Asparagus inferred Over the last 20 years, many scientifi c within the Asparagaceae from plastid petB intron and petD-rpoA papers and reports have been produced The debate over whether the Asparagace- intergenic spacer sequences. They found outlining the establishment, distribution ae contains one genus Asparagus with or evidence supporting a monophyletic ori- and weed status of Asparagus weeds in without subgenera, or up to 16 separate gin of Asparagus and the sub-division of Australia. Differing use of authorship genera has been going for over 200 years. Asparagus into more than three groups. and species names are present in this lit- The taxonomic history of the Asparagace- The Eurasian species of Asparagus formed erature resulting in confusion over which ae is well covered in recent papers. Clif- a monophyletic group, whereas the south- species are actually present. This paper ford and Conran (1987a,b,c) follow Ober- ern African species are potentially para- examines the taxonomic status and au- meyer’s (1983) treatment of the southern phyletic, comprising more than the two thorship of the eight Asparagus species African Asparagaceae species and use the groups (Myrsiphyllum e.g. A. asparagoides naturalized in Australia: A. aethiopicus three genera Asparagus, Myrsiphyllum and (L.) Druce and Protasparagus e.g. A. plu- L., A. africanus Lam., A. asparagoides (L.) Protasparagus for Asparagaceae species in mosus Baker and A. aethiopicus L.) that are Druce, A. declinatus L., A. offi cinalis L., Australia. Malcomber and Sebsebe De- currently recognized in some taxonomic A. plumosus Baker, A. scandens Thunb., missew (1993) determined that the charac- treatments. A. asparagoides is the most dif- and A. virgatus Baker. Asparagus aspara- ters used to separate Protasparagus and As- ferent from all other Asparagus species. goides is found in two forms, one with paragus were insuffi cient and they should The Australian native, A. racemosus is most a widespread distribution and a second both be treated as the sub-genus Asparagus similar to southern African species. form, currently of restricted distribution under the genus Asparagus, with Myrsi- in South Australia and Victoria. There phyllum Willd. also retained as a sub-ge- Current taxonomy of Asparagus are at least nine other Asparagus species nus. Fellingham and Meyer (1995) agreed weeds in Australia in cultivation or present in Australian with Malcomber and Sebsebe Demissew Asparagus aethiopicus L. herbariums, but they are not known to (1993), but found the characters used to SYNONYMS: Protasparagus aethiopicus (L.) be naturalized in bushland. Clarifi cation warrant Myrsiphyllum’s sub-generic status Oberm., Asparagus lanceus Thunb., As- of the nomenclature of Asparagus species to be inconsistent and returned all south- paragus sprengeri Regel and a better understanding of the phy- ern African species to the genus Asparagus CULTIVARS: ‘Sprengeri’ logeny within the family will assist in with no sub-genera. All subsequent papers COMMON NAMES: Sprengers fern, as- the management and policy decisions on on Asparagus have followed Fellingham paragus fern. weed control for each species, for exam- and Meyer’s (1995) treatment. Generally NATIVE DISTRIBUTION: Western and ple, the nomination of various Asparagus the family Asparagaceae is considered to Eastern Cape regions of South Africa. species as target species for biological comprise the one genus Asparagus (Klein- DISTRIBUTION IN AUSTRALIA: Coastal control. jan and Edwards 1999). However, Hemi- northern New South Wales (inc. Lord phylacus, a small Mexican genus, is possi- Howe Island) and southern Queens- Introduction bly included in the family based on recent land, South Australia and Western The family Asparagaceae consists of one evidence from embryology, cytology and Australia. genus, Asparagus, under current taxonom- molecular analyses (Rudall et al. 1998). NOTES: In Green’s (1986) examination of ic revisions and includes approximately the correct name for A. sprengeri, listed as a 120 species mostly of southern African, Evidence from molecular phylogeny synonym of A. densifl orus by Jessop (1966), European and Asian origin. Australia has Three studies have examined the molecu- he found the species to be a cultivar of A. one native species, Asparagus racemosus lar phylogeny of Asparagus species, how- aethiopicus and not A. densifl orus. Green’s Willd., located in far northern Queens- ever two of the studies only included a (1986) assessment was not reflected in land, Northern Territory and northern small number of species. Lee et al. (1997) Obermeyer and Immelman’s (1992) review Western Australia, which has a distribu- examined the phylogenetic relationships of southern African Asparagaceae where tion that extends to north-eastern South of 10 species in the genus Asparagus using they list A. densifl orus as having two cul- Africa, northern Botswana and Namibia. chloroplast DNA. Their research raises the tivars – ‘Sprengeri’ and ‘Myersii’. Follow- All other Asparagaceae in Australia are possibility that dioecy and polyploidy in ing Green (1986), the name A. densifl orus is introduced from southern Africa and Eu- the Asparagus is monophyletic in origin, no longer current in Australian Herbaria, rope (one species, A. offi cinalis L., crop as- but they concede variability in nuclear but the species does exist in cultivation as paragus). At least eight of these species DNA and the inclusion of many more A. densifl orus cv ‘Myersii’. However, the have naturalized in southern and/or cen- Asparagus species is required to further climate-based prediction for A. aethiopicus tral eastern Australian bushlands where understand the taxonomy and evolution and A. densifl orus (Kunth) Jessop indicates they have become environmental weeds. of this genus. Stajner et al. (2002) exam- that Australian material might be more ap- This paper summarizes the taxonomy of ined the nuclear genome size and genetic propriately named A. densifl orus. Asparagus species naturalized in Australia similarity of 10 species of Asparagus based and identifi es issues that require further on their potential application for cross- Asparagus africanus Lam. research. breeding with crop asparagus, Aspara- SYNONYMS: Protasparagus africanus gus offi cinalis. They found the European (Lam.) Oberm. Plant Protection Quarterly Vol.21(3) 2006 129 COMMON NAMES: Asparagus fern NATIVE DISTRIBUTION: Western edge cladodes and branchlets in the one plane, NATIVE DISTRIBUTION: Coastal West- of the Western Cape and Northern and if fruit colour is not important, then ern Cape, Eastern Cape and KwaZulu- Cape regions of South Africa. the specimens are probably A. plumosus. Natal regions of South Africa. DISTRIBUTION IN AUSTRALIA: South- But, the fruit characteristics are usually re- DISTRIBUTION IN AUSTRALIA: Coastal west Western Australia, South-west liable at specifi c level (Jessop 1966) there- South-east Queensland where it is a se- and south-central South Australia (in- fore the confusion in Australian species rious weed (Conran and Forster 1986). cluding Kangaroo Island). still remains. A review of the herbarium NOTES: Obermeyer and Immelman (1992) specimens is required to determine wheth- describe A. africanus as ‘up to 0.6 m or Asparagus offi cinalis L. er there is an error in the description or that more high’. Australian material appears to CULTIVARS: examples ‘Mary Washing- Australia does have both species natural- be much larger implying that it would be ton’, ‘Edulis’, ‘Purple Passion’, ‘Atlas’, ized. worthwhile verifying the identifi cation. It is ‘Jersey Giant’ ‘UC157’. likely that A. africanus, being found in both COMMON NAMES: Asparagus, crop as- Asparagus scandens Thunb. Mediterranean and subtropical climates in paragus, garden asparagus, edible as- SYNONYMS: Myrsiphyllum scandens southern Africa (Obermeyer and Immel- paragus. Thunb. (Oberm.), Asparagus pectinatus man 1992), will need to be re-examined as NATIVE DISTRIBUTION: Southern Eu- Del., Asparagopsis scandens (Thunb.) it may contain more than one species. rope, Northern Africa. Kunth., Asparagus scandens var. defl ex- DISTRIBUTION IN AUSTRALIA: south- us Baker. Asparagus asparagoides (L.) Druce ern Western Australia, southern South COMMON NAMES: Asparagus fern, SYNONYMS: Medeola asparagoides L., Australia, Victoria, New South Wales climbing asparagus fern. Myrsiphyllum asparagoides (L.) Willd., and southern Queensland. NATIVE DISTRIBUTION: Western Cape Asparagus medeoloides Thunb., Elide region of South Africa. asparagoides (L.) Kerguélen, Dracaena Asparagus plumosus Baker DISTRIBUTION IN AUSTRALIA: South- medeoloides L.f., Elachanthera sewelliae SYNONYMS: Protasparagus plumosus west coast of Western Australia, Mount F.Muell., Luzuriaga sewelliae (F.Muell.) (Baker) Oberm. Lofty Ranges and Fleurieu Peninsula K.Krause. COMMON NAMES: Climbing asparagus of South Australia, central coast area of COMMON NAMES: Bridal creeper, bridal fern, ferny asparagus. Victoria and northern Tasmania. veil creeper, fl orist’s smilax, smilax. NATIVE DISTRIBUTION: Eastern South NOTES: A. scandens is a weed in north- NATIVE DISTRIBUTION: Widespread in Africa. ern New Zealand where it is invading the Western Cape, Eastern Cape, Kwa- DISTRIBUTION IN AUSTRALIA: South- the understorey of lowland broad-leaved Zulu-Natal region extending to the west Western Australia, Eyre and and secondary forest (Timmins and Reid Northern Province. Also recorded in Fleurieu Peninsula of South Australia, 2000) Namibia and tropical Africa. north-coast New South Wales (inc. Lord DISTRIBUTION IN AUSTRALIA: Wide- Howe Island) and southern Queens- Asparagus virgatus Baker spread in southern Western Australia, land. SYNONYMS: Protasparagus virgatus (Bak- South Australia, Victoria, northern Tas- NOTES: This species has been referred er) Oberm, mania, sporadic throughout coastal, to as A. setaceus (Kunth) Jessop, but this COMMON NAMES: Asparagus fern central and south-west irrigated re- name may have been misapplied. Clif- NATIVE DISTRIBUTION: Eastern South gions of New South Wales. ford and Conran (1987b) list A. setaceus as Africa extending toward the Northern NOTES: There is currently no taxonomic a synonym of A. plumosus, but they are Province and also Namibia. distinction between the widespread form recognized as two different species by DISTRIBUTION IN AUSTRALIA: Coastal of Asparagus asparagoides and the south- Obermeyer and Immelman (1992) and South-east Queensland. western Cape form as described by Klein- Fellingham and Meyer (1995). Some of jan and Edwards (1999) and discussed in the confusion may stem from the descrip- Other Asparagus species in Coles et al. (2006). Further research on the tion of fruit colour: in Obermeyer and Australia two forms is required to determine if they Immelman’s (1992) description of South There are several other Asparagus species are two different species. African Asparagaceae, they describe the in cultivation in Australia that have not Most recent scientifi c papers incorrectly A. plumosus berry as red and A. setaceus (yet) shown weedy potential. Keighery assign the authorship of A. asparagoides to as black. Conran and Forster (1986) and (1996) lists the following species as com- W. Wight. The correct authorship is A. as- Clifford and Conran (1987b) describe the mon in cultivation in the Perth area; A. paragoides (L.) Druce, as W. Wight, a con- berry of A. plumosus as black and Green densifl orus (Kunth) Jessop, A. falcatus L., A. tributor to the 1909 Century Dictionary (1994) describes A. plumosus berry as red. setaceus (Kunth) Jessop (see notes under and Cyclopaedia, lists A. asparagoides un- Imada et al. (2000) point out that A. setaceus A. plumosus), and A. verticillatus L. Less der the genus Myrsiphyllum in the context has been used extensively in horticultural common species in cultivation include A. of the treatment by Baker (1875), where A. references based on Jessop (1966), but that acutifolius L., A. crispus Lamark, A. tenu- asparagoides is in the sub-genus of Myrsi- the two species involved can be separated ifl orus Lamark and A. virgatus Bak. (Keigh- phyllum (Willd.) Baker. Druce (1914) made by the arrangement of their cladodes: all ery 1996). Randall (2002) lists A. fi licinus the fi rst new combination of A. aspara- in one plane (A. plumosus) or radiating D.Don and A. setaceus as weeds in Aus- goides, based on Medeola asparagoides from in many planes (A. setaceus). In Jessop’s tralia, but we have not located a record of Linnaeus (1753). (1966) revision of the southern African As- establishment. paragaceae where A. plumosus is treated as Asparagus declinatus L. a synonym of A. setaceus, he describes A. Discussion SYNONYMS: Myrsiphyllum declinatum setaceus as having cladodes in one plane In attempting to summarize the current (L.) Oberm., Asparagus crispus Lam., and the berry as red. Imada et al. (2000) understanding of the taxonomy of the Asparagus fl exuosus Thunb., Asparagus conclude that the widespread cultivated eight asparagus weeds in Australia we decumbens Jacq., Asparagopsis decumbens species is A. plumosus and that it is now highlight some areas that require further (Jacq.) Kunth naturalized on three Hawaiian islands. clarifi cation. A clear understanding and COMMON NAMES: Bridal veil, bridal Both Green (1994) and Clifford and Con- application of the names of species is im- veil creeper, bridal creeper. ran (1987b) describe A. plumosus as having portant for any future policy decisions 130 Plant Protection Quarterly Vol.21(3) 2006 including the application of noxious weed Australian Herbarium for their comments Western Australia. Plant Protection legislation and the nomination of species on the manuscript. Quarterly 11, 49-50. weeds (excluding A. officinalis) as tar- Kleinjan, C.A. and Edwards, P.B. (1999). gets for biological control. Of particular References A reappraisal of the identifi cation and relevance is the use of A. aethiopicus/A. Baker, J.G. (1875). Revision of the genera distribution of Asparagus asparagoides in densifl orus and A. plumosus/A. setaceus as and species of Asparagaceae. Journal of southern Africa. Southern African Jour- synonyms for each other between the Aus- the Linnean Society, Botany 14, 508-632. nal of Botany 65, 23-31. tralian States. In the case of AA.. plumosusplumosus/A. Clifford, H.T. and Conran, J.G. (1987a). Lee, Y., Kanno, A. and Kameya, T. (1997). setaceus, further examination of herbarium 2. Asparagus. Flora of Australia 45, 159. Phylogenetic relationships in the genus specimens is required to determine if both Clifford, H.T. and Conran, J.G. (1987b). Asparagus based on the restriction en- species are naturalized in Australia. The 3. Protasparagus. Flora of Australia 45, zyme analysis of the chloroplast DNA. Western Cape form of ‘Asparagus aspara- 159-63. Breeding Science 47, 375-8. goides’ (Coles et al. 2006), needs to be stud- Clifford, H.T. and Conran, J.G. (1987c). Linnaeus, C. (1753). Species plantarum: a ied and named so it can be identifi ed and 4. Myrsiphyllum. Flora of Australia 45, facsimile of the fi rst edition 1753 volume targeted for control. Finally, the identifi ca- 163-5. 1. The Ray Society, London, 560 pp. tion of Australian material as A. africanus Coles, R.B., Willing, K.L., Conran, J.G. and Malcomber, S.T. and Sebsebe Demissew also needs verifi cation, but, along with the Gannaway, D. (2006). The identifi ca- (1993). The status of Protasparagus and issues mentioned above, this may depend tion and distribution of Western Cape Myrsiphyllum in the Asparagaceae. Kew on a better understanding of the classifi - form of bridal creeper (Asparagus as- Bulletin 48, 63-78. cation of Asparagus species in southern paragoides) in the south east of South Obermeyer, A.A. (1983). Protasparagus Africa. Australia and Western Victoria. Plant Oberm. nom. nov.: new combinations. Overall, the understanding of the phyl- Protection Quarterly 21, 104-8. South African Journal of Botany 2, 243-4. ogeny of Asparagaceae species is improv- Conran, J.G. and Forster, P.I. (1986). Obermeyer, A.A. and Immelman, K.L. ing (Fukuda et al. 2005) although so far Protasparagus africanus (Asparagaceae) a (1992). Protasparagus. Flora of Southern 24 out of the 100 to 300 species have been serious weed for south-eastern Queens- Africa 5, 11-70. studied. The phylogeny is important be- land. Austrobaileya 2, 300-4. Randall, R.P. (2002). ‘A global compen- cause it will provide hypotheses to test Druce, G.C. (1914). Report of the Botanical dium of weed’s. (R.G. and F.J. Richard- the host range of potential biological con- Exchange Club of the British Isles 1913, son, Melbourne). trol agents and to provide explanation of 3, 414. Rudall, P.J., Engleman, E.M., Hanson, L. host range patterns. For example, future Fellingham, A.C. and Meyer, N.L. (1995). and Chase, M.W. (1998). Embryology, biological control of Asparagaceae species New combinations and a complete list cytology and systematics of Hemiphy- will largely be determined by how closely of Asparagus species in southern Africa lacus, Asparagus and Anemarrhena (As- related the weed is to the crop species, A. (Asparagaceae). Bothalia 25, 205-9. paragales). Plant Systematics and Evolu- offi cinalis and to the native A. racemosus. Fukuda, T., Ashizawa, H., Suzuki, R., tion 211, 181-99. Finally, researchers and people with an in- Ochiai, T., Nakamura, T., Kanno, A., Scott, J.K. and Batchelor, K.L. (2006). terest in weeds should lodge specimens of Kameya, T. and Yokoyama, J. (2005). Climate-based prediction of potential asparagus weeds with offi cial herbariums Molecular phylogeny of the genus As- distribution of introduced Asparagus to both improve our knowledge of distri- paragus (Asparagaceae) inferred from species in Australia. Plant Protection butions, the range of variability present in plastid petB intron and petD-rpoA inter- Quarterly 21, 91-8. Australia including new naturalizations, genic spacer sequences. Plant Species Stajner, N., Bohanec, B. and Javornik, B. and to provide the specimens needed to Biology 20, 121-32. (2002). Genetic variability of economi- elucidate the questions mentioned in this Green, P.S. (1986). The correct name for cally important Asparagus species as paper. Asparagus sprengeri. The Plantsman 7, revealed by genome size analysis and 249-50. rDNA ITS polymorphisms. Plant Sci- Acknowledgments Green, P.S. (1994). Liliaceae. Flora of Aus- ence 162, 931-7. We thank Carien Kleinjan, University of tralia 49, 502-13. Timmins, S.M. and Reid, V. (2000). Climb- Cape Town, South Africa and Karen Wil- Imada, C.T., Staples, G.W. and Herbst, D.R. ing asparagus, Asparagus scandens son, New South Wales Herbarium for dis- (2000). New Hawaiian plant records for Thunb.: a South African in your forest cussions on Asparagus species identifi ca- 1999. Bishop Museum Occasional Papers patch. Austral Ecology 25, 533-8. tion. Thanks also to John Conran, Univer- 63, 9-16. Wight, W.F. (1909). (No title). In ‘The cen- sity of Adelaide, South Australia, David Jessop, J.P. (1966). The genus Asparagus in tury dictionary and cyclopedia: a work Cooke, Department of Water, Land and Southern Africa. Bothalia 9, 31-96. of universal reference in all departments Biodiversity Conservation, South Austral- Keighery, G. (1996). Native, natural- of knowledge’, ed. B.E. Smith, XI p. 845. ia and Terry MacFarlane from the Western ized and cultivated Asparagaceae in (The Times Book Club, London).