Madrono, Vol. 55, No. 3, pp. 244^247, 2008 NOTES ON TWO SOUTHERN AFRICAN ARCTOTIS SPECIES (ARCTOTIDEAE: ASTERACEAE) GROWING IN CALIFORNIA Alison M. Mahoney Department of Biology, Minnesota State University - Mankato, MN 242 Trafton Science Center S, Mankato, 56001 [email protected] Robert McKenzie J. Molecular Ecology and Systematics Group, Department of Botany, Rhodes University, P.O. Box 94, Grahamstown, 6140, South Africa Abstract A literature review and determination of specimens performed in conjunction with treatment preparations for the Flora ofNorth AmericaNorth ofMexico and the secondedition ofTheJepson Manualindicatesthat thenamesinusefortwoArctotisspecies(Arctotideae: Asteraceae)occurringin California needupdating. Venidiumfastuosum (Jacq.) Stapf, a rareescapefromcultivation, shouldbe Arctotisfastuosa Jacq. (PI. hort. schoenbr. 2: 20, pi. 166; 1797) and A. venusta Norl. (Bot. Not. 118: 406-7; 1965) is the correct name for a naturalized species previously determined as A. stoechadifolia P.J. Bergius. Key Words: Arctotideae, Arctotis, Asteraceae, naturalization, Venidium. The only members oftribe Arctotideae (Aster- Venidiumfastuosum aceae) with established populations in North America occur in Cahfornia and New Mexico The genus Arctotis is characterized principally (Mahoney 2006). All are southernAfrican species by achenes with three ± well-developed abaxial introduced through horticulture. Among them, wings that create one or two distinct furrows or only one species is potentially invasive; the others s"ceagvrietgiaetse"d(fMrcoKmenArzciteoteitsabl.y2L0e0s5s)i.ngVe(n18i3d1i,um18w3a2s) range from occasional to naturalized garden escapes with limited distributions. Munz and based on the less well-developed achene wings, the absence or extreme reduction of pappus Keck (1973) and McClintock (1993) treated four scales, and differences in achene pubescence. species: Gazania linearis (Thunb.) Druce [G. Beauverd (1915) and Lewin (1922) found the longiscapa DC], Arctotheca calendula (L.) Le- distinguishing characters used by Lessing unten- vyns, Venidium fastuosum (Jacq.) Stapf, and able and transferred Venidium species into Arctotis stoechadifolia P.J. Bergius. Arctotis, but Stapf (1926) disagreed and trans- NaturalizedpopulationsofG. linearis(treasure- ferred A. fastuosa Jacq. to Venidium. Use of the flower) occur along roadsides, especially in urban name V. fastuosum has persisted in U.S. floral coastal areas of Santa Barbara and Los Angeles treatments (Munz and Keck 1973; Liberty Hyde Counties. EntitiestreatedpreviouslyasArctotheca Bailey Hortorium 1976; McClintock 1993; Quat- calendula (capeweed) (Munz and Keck 1973; trocchi 2000; Calflora 2007; USDA 2007). McClintock 1993) actually consist oftwo distinct A morphological study of achenes and initial species. Arctotheca prostrata (Salis.) Britten is a molecular analyses of subtribe Arctotidinae sterile perennial that spreads aggressively by (McKenzie et al. 2005, 2006; Funk et al. 2007) prostrate stems along roadsides and in other support Beauverd's and Lewin's observations for disturbed sites in the North Coast (NCo), South some of the species previously placed in Veni- Coast (SCo), Central West (CW), and Western dium. Results from a larger molecular study of Transverse Ranges (WTR) floristic province Arctotidinae (McKenzie and Barker 2008) indi- subregions ofCalifornia. Arctotheca calendula, a cate that although Arctotiss.l. is polyphyleticand fertile annual classified as invasive by the Cali- needs redefining, California's Arctotis species will fornia Exotic Pest Plant Council (Brossard et al. retain their names. Based on these new data, the 2000), occursinafewcoastalanddisturbedurban name Arctotisfastuosa Jacq. should be adopted habitats in the NCo, Central Coast (Ceo), and for the species previously known as Venidium Outer South Coast Ranges (SCoRO) subregions. fastuosum. The present paper is confined to updating Arctotis fastuosa is a hirsute to lanate, tap- names and identities of Venidium fastuosum rooted annual with heads up to 10 cm in (monarch-of-the-veld) and Arctotis stoechadifolia diameter. Its rays are bright orange to yellow (a (blue-eyed African daisy). white-rayed form is in cultivation), usually 2008] MAHONEY AND MCKENZIE: TWO ARCTOTIS SPECIES IN CALIFORNIA 245 marked basally with a purple-black band; its flowering shoots arise. Its ray florets are white or discs areyellowish-brown. The species is native to pale yellow; its discs are black. The silver-grey the semi-arid, winter-rainfall Namaqualand re- leaves have a dense, tightly appressed woolly gion of South Africa. Stapf(1926) referred to A. tomentum on both the upper and lower surfaces. fastuosa as a weed of cornfields in its native The outer involucral bracts have a rigid base and range, where it is now common on roadsides, a 4—7 mm long, acuminate, linear-cylindrical but in California it very rarely escapes from apical appendage. The obovoid-obconical cultivation and does not appear to have natural- achenes are densely tomentose and bear strongly ized. We are not aware of the species being a incurved, ± entire lateral wings that partially problematic weed in any country. McKenzie conceal the abaxial 'cavities.' Arctotis venusta is et al. (2005) found that herbarium specimens an erect, tap-rooted, summer-flowering annual. determined as A. fastuosa can be divided into Its ray florets are white with a narrow yellow at least three distinct entities based on achene band at the base of the limb; its discs are a morphology. We have been able to examine distinctive greyish-purple. The leaves are usually achenes from two CaUfornian specimens {Fuller more thinly and loosely tomentose. The outer 8191, CDA; R. Whitaker s.n., RSA); in both, involucral bracts have a soft base and a short, 1- mm achenes lack pappi, pubescence, and basal tufts 3 long appendage with a rounded or obtuse of hairs and are readily referable to typical A. apex. The oblong-obconical achenes are ± fastuosa. glabrous to sparsely tomentose on the abaxial Specimens examined. USA, CA, Riverside Co., and tangential surfaces and the oblong-obovate Thermal, estabhshing itself, 10 March 1949, cavities are obvious. Herbarium specimenscan be Whitaker s.n. (RSA), 5.2 mi S of Beaumont, more difficult to distinguish due to faded fiorets roadside sand, 17 February 1960, Fuller 3609 and missing lower-stem or below-ground parts, (CDA); San Bernardino Co., San Bernardino, but the differences in leafpubescence, involucral- NE of Central Ave. & 3'^'* St, Gate 5, Norden bract and achene features allow discrimination of AFB, waste ground, 11 April 1962, Fuller 8191 the two species. (CAS, CDA). Our determination of CaUfornian specimens from horticulture and naturally-occurring popu- Arctotis stoechadifolia and A. venusta lations previously determined as A. stoechadifolia During a 1939^0 study ofArctotis specimens, confirms that they belong to A. venusta. No Norlindh (1964) found that in the last revision of specimens of A. stoechadifolia have been ob- Arctotidinae, Lewin (1922) had misinterpreted A. served. Norlindh (1964) noted that A. venusta is stoechadifolia P.J. Bergius, a prostrate, mat- weedy even in its native range, often occurring in forming perennial occurring in open sandy, cultivated fields or along roadsides, while A. seasonally wet areas and stabilized dunes in the stoechadifolia does not readily tolerate competi- southwestern Cape. Lewin applied this name to tion so that some populations have disappeared an erect, tap-rooted annual occurring in a wider due to urban sprawl and the planting oferosion- range ofhabitats and with a broader distribution preventing trees and shrubs. Wells et al. (1986) in inland southern Africa (Norlindh 1964). The classified A. venusta as a ruderal, agrestal and native ranges of the two entities do not overlap. pastoral weed in South Africa. Stated undesirable Although Norlindh (1965) subsequently de- characteristics include: that it may be poisonous scribed and named the annual species A. venusta to stock, it taints milk, and is a crop-seed Norl., Lewin's interpretation has been adopted contaminant. Arctotis venusta is predicted as widely. Arctotis venusta is a popular horticultural highly likely to become a weed, and further- plant and is often grown under the name A. more to become an agricultural weed, in Aus- stoechadifolia, A. stoechadifolia var. grandis tralia (Scott and Panetta 1993). It appears (Thunb.) Less, or A. grandis Thunb. Sometimes that A. venusta has been cultivated in California the names A. stoechadifolia and A. venusta are for at least 150 years but in that time has not treated as synonyms (Brickell and Zuk 1996). naturalized extensively. However, all but one of Mahoney's (2006) FNA treatment maintained the Californian specimens of A. venusta we the name A. stoechadifolia but commented on its examined were collected before 1971; whether A probable misuse. full investigation subsequent this reflects its rarityora lull in activecollectingis to preparation of that treatment and availabiUty unclear. It is interesting that naturalized Califor- of further data have clarified the identity of the nian populations of A. venusta occur in coastal, Californian specimens. sandy habitats that might be favored by A. Arctotis stoechadifolia and A. venusta are stoechadifolia, even though A. venusta is an morphologically distinct species (Norlindh inland species occurring in rangeland habitats 1964). Fresh plants cannot be confused. Arctotis far from the coast in its native range in southern stoechadifolia is a mat-forming perennial that Africa. produces long (up to ± 1 m), adventitiously The true Arctotis stoechadifolia has become rooting, prostrate stems from which the erect naturalized along parts of the coastline of MADRONO 246 [Vol. 55 Australia (Jeanes 1999; Rippey and Rowland Brickell, C. and J. D. Zuk (eds.) 1996. American 2004; Barkeret al. 2005) due to its popularity as a Horticultural Society: A-Z encyclopedia ofgarden garden ornamental plant and its use as a dune plants. Dorling Kindersley Publishing, New York, NY. ostpaebinliszietre.s iItn tihnehapbritostecttheedfhoirned--dduunneetahrreoa,ugahs tion BrosHsoasrHdO,VSCK.YC(,eds.J). 2M00.0.RIannvdasailvle,plaanntsdofM.CalCi.- its indigenous range in South Africa. The species fornia's wildlands. University of California Press, has been assessed to be a minor problem weed in Berkeley, CA. natural ecosystems warranting control at four or Calflora, 2007. Information onCaliforniaplants for more locations within a state or territory in education, research and conservation [web appli- Australia (Groves et al. 2003). In parts of South cation]. The Calflora Database, Berkeley, CA. Australia, it has a high Weed Risk Assessment Available at: http://www.calflora.org. Accessed rating, as the dense mats smother smaller March 03, 2007. indigenous plants by shading and competition Cordingley, S. p. and C. E. Petherick. 2005. for resources and can alter dune shape (Cording- VaengdetWaetsiotnBmeaancahgeDmuennet pRleasnerfvoer.thCeitHyenolfeyChSaorultehs ley and Petherick, 2005). Therefore, if A. Sturt, Adelaide, Australia. stoechadifolia is cultivated in California we Funk, V. A., R. Chan, and A. Holland. 2007. recommend assessment of its weed risk poten- Cymbonotus (Compositae: Arctotideae, Arctotidi- tial and vigilance for naturalized plants to ensure nae): an endemic Australian genus embedded in a it does not become a problem species in southern African clade. Botanical Journal of the California. Linnean Society 153:1-8. Specimens examined. USA, CA, Los Angeles Groves, R. H., J. R. Hosking, G. N. Batianoff, Co., Glendora, Alosta Ave. at Glendora Ave., D. A. CooKE, I. D. CowiE, R. W. Johnson, G. J. 750 ft., dry roadside, 24 June 1933, Wheeler 1889 KMeoiEgRhKeErRyK,,B.R.J.p.LeRpasncdhia,lla,. AA.. CM.itRcohzeelfle,ldMs., (CDA, RSA, UCR); [county name illegible, N. G. Walsh, and B. M. Waterhouse. 2003. probably Los Angeles], Ocean Park, sand lots Weed categories for natural and agricultural near beach, 2 August 1934, Cohen [?] 575 (RSA); ecosystem management. Bureau ofRural Sciences, Orange Co., 2 mi NE ofHuntington Beach near Canberra, Australia. N end of Bolsa Chica Salt Marsh, disturbed Jeanes, J. A. 1999. Asteraceae. Vol. 4, Pp. 652-984 in roadside, infrequent escape, 25 July 1970, Hen- Walsh N. G. and T. J. Entwistle (eds.). Flora of rickson 5091 (RSA), Newport Bay, 10 ft., sandy Victoria. Inkata Press, Melbourne, Australia. flat just back of high tide, 21 October 1933, Lessing, C. F. 1831. Synanthereae Rich. Pp. 83-170 in MWahreieal,erN22s2id4e(oRfSWA);JoSnaenstaStBEarobfarSaCCuor.v,erSa[nsitca. ctLo.hrtieCan.draAol.ma(vneodzsno.f).CfihDaaenmaipslosabnosteirasvnaditnisDe.xdpiesFds.iertLei.roen.veoLnsipneSnccauhellaae,-- probably Curryer] St., naturalized in waste 6:76-170. ground, 21 May 1968, Fuller 17065 (CDA), Santa . 1832. Synopsis generum compositarum ear- Maria, 100 E Main, vacant lot, 22 September umque dispositionis novae tentamen, monogra- 1965, Jones & Allen s.n. (CDA), Lauro Canyon phiismultarumcapensiuminterjectis. Dunckerand Dam Project, roadside, 30 May 1952, Pollards.n. Humblot, Berlin, Germany. (CDA), Hwy between Santa Maria and Orcutt, Lewin, K. 1922. Systematische Gliederung und geo- sandy soil along highway, 22 November 1960, graphische Verbreitung der Arctotideae-Arctotidi- Smith 6332 (RSA, SBBG, UCR); Ventura Co., nae. Repertorium specierum novarum regni veget- Ojai, Canada St., 12 July 1946, Pollard s.n. Libearbitlyis,HyBediehefBtaeil1e1:y1-H7o5.rtorium. 1976. Hortus III. (CAS). MacMillan Publishing Co., New York, NY. Mahoney, a. M. 2006. Arctotis. Vol. 19, Pp. 198-199 Acknowledgments in Flora of North America Editorial Committee We thank David Keil (OBI) and Andrew Sanders (ed.). Flora of North America North of Mexico. (UCR) for insights and assistance; the curators ofthe Oxford University Press, New York, NY. CAS, CDA, RSA, SBBG, SD and UC herbaria for the McClintock, E. 1993. Arctotis, Pp. 200; Venidium, p. loan of, or access to, specimens; and the National 356 in J. C. Hickman (ed.). The Jepson manual: Research Foundation of South Africa for a Postdoc- higherplantsofCalifornia. UniversityofCalifornia toral Fellowship to R. J. McK. McKePrnezsis,e,BeRr.keJl.eya,nCdAN.. p. Barker. 2008. Radiation Literature Cited of southern African daisies: Biogeographic infer- ences forsubtribeArctotidinae (Asteraceae, Arcto- Barker, W. R., R. M. Barker, J. P. Jepson, and tideae). Molecular Phylogenetics and Evolution H. P. VONOW (eds.) 2005. Census of South 49:1-16. Australian vascular plants. 5"' ed. Journal of the , E. M. MuLLER, A. K. W. Skinner, P. O. Adelaide Botanic Gardens Supplement 1. Botanic Karis, and N. p. Barker. 2006. 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