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Status of Introduced Plants in Southern Arizona Parks Dimorphotheca sinuata D.C.
William L. Halvorson, Principal Investigator U.S. Geological Survey / Southwest Biological Science Center Table of Contents:
Dimorphotheca sinuata D.C. 3
African daisy, cape marigold, sun marigold, Star of the Veldt, glandular cape marigold.3 synonymous names of the species:. 3
species taxonomy. 3
similar native or non-native species that could confuse identification. 5
biology. 5
ecology. 6
climatic requirements and limitations:.6 why it has the potential to do well as an exotic:.7 effect on natural processes/description of the threat. 8
known general distribution. 8
National Park Service, southern Arizona group:.9 Montezuma Castle National Monument and Montezuma Well unit.9 control methods and management strategies. 10
contacts or technical specialists. 10
bibliography. 11
additional sources and websites. 14
Dimorphotheca sinuata D.C.
African daisy, cape marigold, sun marigold, Star of the Veldt,
glandular cape marigold

family: Asteraceae

synonymous names of the species:
the first name in each species list is the current and synonymous name used by the name in bold type occurring within each species list indicates the plant name used within these documents, which is also the name provided in the southern Arizona NPS exotics database 'soaraz~1.xls' (Holden 1996). Osteospermum sinuatum (D.C.) Norl. Dimorphotheca sinuata D.C.
other names found: Dimorphotheca aurantiaca species taxonomy

Dimorphotheca sinuata D.C., African daisy: From Hickman (1993), Munz (1974),
(A glossary is provided at the end of this section for the plant terminology used in this section.) life strategy: an annual, herbaceous plant. Reproduces from seeds. 2n=18.
structure: an erect, usually branched, annual herbaceous plant; up to 12 in. (3 dm) tall.
roots: small taproot with fibrous branching roots.
stems: stems 4-12 in. (10-30 cm) long.
branching: stems simple to sparingly branched from base.
stipules: no stipules.
leaves: leaves alternate along stem. Leaves less than 4 in. (10 cm) long, oblong to
oblanceolate, entire to coarsely sinuate-dentate; lower leaves tapered to petiole-like base; upper leaves sessile, smaller, sometimes linear. Herbage glandular-pubescent. inflorescence: composite heads solitary on terminal, long peduncles, 1.2-2.8 in. (3-7 cm)
phyllaries: involucre broadly campanulate; phyllaries in one series, linear-lanceolate,
0.4-0.6 in. (10-15 mm) long, acuminate, narrowly scarious-margined. corolla/petals: receptacle flat or nearly so, naked. Ray flowers fewer than 15, pistillate,
corollas yellow to orange, sometimes violet at base or apex, ligules 0.8-1 in. (2-2.5 cm) long. Disc flowers many, perfect, corolla 0.2 in. (4.5-5.5 mm) long, corolla with very short tube and long throat, yellow to orange, often purple tipped. No pappus. gynoecium: ray flowers with style divided into two long stigmatic branches. Disc
flowers with style divided into two short stigmatic branches. androecium: anthers with ovate or triangular-ovate tips, sagittate at base, short-tailed.
fruit: achenes of ray flowers 3-angled to subterete, 0.2 in. (4-5 mm) long, usually
wrinkled or tuberculate. Achenes of disc flowers flattened, 0.1-0.3 in. (6-7 mm) long, obovate, smooth, winged or thickened margins, no pappus. taxonomic glossary (Harris and Harris 1997):
acuminate: tapering to a sharp point
campanulate: bell-shaped
dentate: toothed along the margin, the teeth directed outward rather than forward (serrate)
lanceolate: much longer than wide; with the widest point below the middle
oblanceolate: inversely lanceolate, with attachment at the narrow end
obovate: inversely ovate, with attachment at narrow end
obtuse: blunt or rounded at the apex
ovate: egg-shaped outline, with attachment at broad end
peduncle: the stalk of a solitary flower or of an inflorescence
perfect: having both male and female reproductive organs
petiole: leaf stalk
pubescent: covered with short, soft hairs
sagittate: arrowhead-shaped with the basal lobes directed downwards
scarious: thin, dry, and membranous in texture, not green
sessile: attached directly, without a stalk
sinuate: having a strongly wavy margin
sub-: meaning under, slightly, somewhat, or almost -
terete: round, in cross section
tubercle: a small tuber-like swelling or projection
tuberculate: having tubercles
image of plant

obtained at CalPhotos, Berkeley Digital Library Project: obtained at CalPhotos, Berkeley Digital Library Project: http://elib.cs.berkeley.edu/cgi/calflora_query?one=T&special=photo& http://elib.cs.berkeley.edu/cgi/calflora_query?one=T&special=phot obtained at CalPhotos, Berkeley Digital Library Project: Dimorphotheca sinuata. Website: http://elib.cs.berkeley.edu/cgi/calflora_query?one=T &special=photo&where-calrecnum=2733 similar native or non-native species that could confuse identification

Although there are many plants in the desert that are early blooming and daisy-like, African
daisies/Cape marigolds are one of the earliest bloomers, salmon colored to yellow to orange, and seemingly have no other species that it would be readily misidentified with during its biology

growth and reproductive strategy:
Dimorphotheca sinuata germinates following sufficient rains in the fall and winter and into spring (van Rooyen et al. 1992a, 1992b). Over the winter the plant grows vegetatively, adding biomass. Flower initiation occurs after a drop in temperature during the winter (vernalization) with flowering usually occurring in the spring after temperatures have begun to rise (van Rooyen et al. 1991, Steyn et al. 1996). Typical of annual plant strategies, Dimorphotheca sinuata completes its life cycle during favorable environmental conditions and the species survives stressful environmental conditions (i.e. drought and heat) as a seed (van Rooyen et al. 1992b). This species showed a dependence on insect pollination for cross-pollination and successful seed set. Self-compatibility systems were operative, but self-pollination is avoided in this plant by protrandry (maturity of male reproductive system before maturity of female reproductive system) (Ueckermann and van Rooyen 2000). Dimorphotheca sinuata produces dimorphic fruits (two different achenes/seeds). These achenes/seeds are morphologically and anatomically different, with each having different responses to environmental conditions, ensuring species' survival in stressful climates (Beneke et al. 1992a, 1992b, 1993a, 1993b, 1993c). no sources found for potential number of seeds/plant. Beneke et al. (1993a) reported that disc achene: ray achene ratios changed for plants growing in different localities Seeds tend to drop from the plant as soon as they are mature (Knowles et al. 1965). Achenes of the disc flowers are winged, having better dispersal mechanisms than achenes of the ray flowers, which are non-winged (Beneke et al. 1993b). Bass (1977) reports on laboratory germination and longevity tests of crop seeds of Dimorphotheca sinuata. He found when seeds of various ages were germinated at 59°F (15°C) in germination blotters, germination ranged between 54-73% for seeds less than a year old, 71-84% after 6 years of storage, 71-84% after 11 years, 71-80% after 16 years. Seeds were stored at 41°F (5°C) and at 40% relative humidity. (He reported that reduced germination in the trials of the seeds less than a year old may have occurred because those achenes were not processed to remove non-filled fruit as the achenes of older stock were, or the seeds were potentially dormant). Bass also referred to a past paper, Bass et al. (1967), in which storage conditions were tested. It was found that storage environment had an impact on Dimorphotheca sinuata seed longevity. Some horticultural sources report that seeds from this plant have ecology

origin and history of introduction:
Dimorphotheca sinuata is native to Africa (Angola, Namibia, the Cape Province of South Africa (Namaqualand)) (GRIN 2000, Hickman 1993, Munz 1974). Dimorphotheca sinuata has been cultivated in Arizona since the 1940's, and has naturalized in various locations since the 1970's (Earle 1973 in Burgess et al. 1991). At the desert laboratory in Tucson the first collection was made in 1978 by Turner and Goldberg; in the 1991 report, Dimorphotheca sinuata was reported to be invading edges of the property; it spreads along washes (Burgess et al. 1991). Hickman (1993) states that it is an escape from cultivation, and establishes along roadsides and in disturbed places in California. Dimorphotheca sinuata seeds have been identified for quality seed oil; the plant has been investigated for its crop In its native area: Dimorphotheca sinuata's native region is characterized by hot, dry summers, with sparse and erratic rainfall occurring primarily in the winter months (Schulze 1965 in Rösch et al. 1997). On the North American continent: In the Sonoran Desert, Tucson, Arizona, Burgess et al. (1991) reports that it spreads along washes. In California, it occurs under 3280 ft. elevation, along roadsides and disturbed areas (Hickman 1993). Dimorphotheca sinuata likes to grow in full sun; its flowers will close in shade. The plants grow over a wide range of temperatures and photoperiods, provided there is sufficient moisture; although it is a drought tolerant plant. Dimorphotheca sinuata produces two different types of achenes within each inflorescence; one type from the disc flowers, another from the ray flowers. The germination requirements differ for each type. The disc achenes/seeds are winged, smooth, and larger; the ray achenes/seeds are smaller, not winged, with a wrinkled to tuberculate pericarp which is thicker and tougher. Optimum germination requirements for both disc and ray achenes was found to be between 45-72°F (7- 22°C) in light, although the disc achenes/seeds germinated at a rate of 95.5% and the ray achenes/seeds germinated at a rate of 3% (Beneke et al. 1993b). de Villiers et al. (1994) found that the highest germination percentages occurred at 68°F (20°C). Germination occurs when there is sufficient moisture available in the fall and winter months, although it also can germinate later, in spring. High temperatures reduced germination; this response was suggested as an adaptation to limit germination after rain in the warm, dry season (Beneke et al. 1993b). The differences between the two types of achenes/seeds are due to anatomical and chemical qualities in the pericarp. The ray achenes/seeds' pericarp not only has extensive sclerenchymous zones and tannins present (mechanical and chemical dormancies), but also a mucilaginous epidermis (Beneke et al. 1992b). Beneke et al. (1993b) reported that the disc achenes/seeds germinated more readily over a wide range of temperatures, in light and dark conditions, and in greater percentages when the conditions were favorable (Beneke et al. 1992b, Beneke et al. 1992a). The mechanical and chemical components of the ray achenes/seeds delay germination during unfavorable conditions (Beneke et al. 1992a, 1992b). Then, when the seeds germinate, the mucilaginous epidermis may contribute to water economy, keeping the achene moist for prolonged periods of time to endure germination (Beneke et al. 1992b). Fruit polymorphism represents a survival strategy; it assures species expansion under favorable conditions and survival under unfavorable conditions Dimorphotheca sinuata likes well-drained soils, and will tolerate dry soils (Michigan there is no information concerning this plant's abilities to compete in non-native situations. Many studies were available describing its competitive abilities intraspecifically, or interspecifically within its native environment (see bibliography); none were available for the North American continent. In Knowles et al. (1965), it was reported that in some of the crop trials on this continent (Georgia, Iowa, Nebraska, North Carolina, Texas, Washington), the plants were plagued by beetles, nematodes, or various fungi, preventing successful growth and/or maturity, but in a number of places (California, Colorado, Kansas, Maryland, Oregon) the crop was more or less successful. why it has the potential to do well as an exotic: Dimorphotheca sinuata is native to a relatively similar stressful, arid environment as the Sonoran Desert. It likes desert soils and environments, and is drought tolerant. As an annual forb, its life cycle is synchronized with the climate, actively growing only when favorable conditions are presented, escaping unfavorable conditions (e.g. high temperatures, water stress/drought) as a seed (van Rooyen et al. 1992b). In its native environment, rains are unpredictable, and this ephemeral plant has adapted by acquiring highly flexible (plastic) responses in lifespace, size, biomass allocation (van Rooyen et al. 1992a, 1992b); rapid establishment, growth, and reproduction are advantages in stressful environments. In addition to these responses, its dimorphism of achenes/seeds lends to an enhanced survivability in variable environments. Further research also showed that the plants that grow from these differing seeds have their own differing capabilities; plants cultivated from the disc achenes/seeds produced more achenes/seeds than those from the ray achenes/seeds. Yet, when under drought conditions, plants from the ray achenes/seeds produced more achenes/seeds (Beneke et al. 1993a). It was also shown that late-sown disc achenes/seeds produced fewer achenes/seeds than if early-sown, while plants grown from the ray achenes/seeds were not affected by sowing date effect on natural processes/description of the threat

no information or sources found: this plant has been encouraged/promoted as an ornamental
for low water use situations and for its spectacular displays (as in its native south Africa); until recently, it hasn't been targeted as an invasive non-native species; information is minimal concerning its status as an invasive plant. known general distribution

United States:
California, Oregon (USDA,NRCS, The PLANTS database 2001: Map available at Website: http://plants.usda.gov/plants/ ; then enter the common or scientific name), Arizona (Holden 1996, Rondeau et al. 1996). Arizona, by county:
Rondeau et al. (1996) lists Dimorphotheca sinuata (African daisy) in Saguaro National Park, Pima County. Niehaus et al. (1984) states that it occurs in southwest Arizona in
National Park Service, southern Arizona group:

Casa Grande Ruins National Monument
source listing species' presence in park: source listing species' presence in park: source listing species' presence in park: source listing species' presence in park: Montezuma Castle National Monument and Montezuma Well unit source listing species' presence in park: source listing species' presence in park: source listing species' presence in park: Rondeau, R., and R. Van Devender. 1992. Floristic survey of the proposed Wildhorse Canyon Protected Natural Area: Final Report. National Park Service, Funding #1443 Rondeau, R., T.R. Van Devender, C.D. Bertelsen, P. Jenkins, R.K. Wilson, M.A. Dimmitt. 1996. Annotated flora and vegetation of the Tucson Mountains, Pima County, Arizona. Desert Plants 12(2):3-46. source listing species' presence in park: source listing species' presence in park: source listing species' presence in park:
Weeds in the West Project
While completing distribution mapping between Spring 1999 through Spring 2001 for the
USGS Weeds in the West project in the southern Arizona National Park Service management units, Dimorphotheca sinuata (African daisy) was found in the following parks Note: A few plants were observed along a dirt roadside in Saguaro National Park during the spring of 2001 during a non-work related hike in the Tucson control methods and management strategies

no information or sources found.
contacts or technical specialists

Dr. Francis E. Northam (Ed Northam)
Noxious Weed Coordinator, Plant Services Division Phone: (602) 542-3309: FAX: (602) 542-1004 Ed works state-wide primarily with noxious agricultural weeds, yet has also done some work to get non-native invasive plants listed that impact Arizona's natural He indicated he would provide, as requested, information regarding: weed biology control/management of weeds
Dr. John H. Brock
Coordinator of Sustainable Technologies, Agribusiness and Resources (STAR) Research Phone: (480) 727-1240; FAX (480) 727-1961 e-mail: invasive plant work (including control treatments) in essentially all the major vegetation types in Arizona, except the highest elevation types like mixed
April Fletcher, Arizona Interagency Weed Action Group
April works region-wide with on-the-ground folks. Arizona Interagency Weed Action Group (IWAG) is an ad-hoc group; working on specific projects identified as species of concern by the group. IWAG consists of invasive weed folks from state and Federal acquainted with control methods for numerous species she knows many professionals who are doing control work, so, when she can't supply an answer, she can usually provide contacts who can.
Jim Horsley, Southwest Vegetation Management Association
Jim indicated at ADOT they manage and control a number of native and non-native invasive species. Their experience includes Centaurea solstitialis (Yellow) and Centaurea melitensis (Malta) star thistle, Onopordum acanthium (Scotch), Carduus nutans (Musk), and Cirsium vulgare (Bull) thistle, Acroptilon repens (Russian), Centaurea biebersteinii / Centaurea maculosa (spotted), and Centaurea diffusa (diffuse) knapweed, Alhagi maurorum (Camelthorn), Halogeton glomeratus (Halogeton), Salsola sp. (Russian thistle, tumbleweed), Linaria damatica (Dalmation toadflax), Cardaria draba (Hoary cress), Tribulus terrestris (Puncture vine), Cenchrus sp. (sandbur), Convolvulus arvensis (Field bindweed), Sorghum halepense (Johnsongrass), Pennisetum ciliare (Buffelgrass), Pennisetum setaceum (Fountain grass), several mustards, Verbascum sp. (mullein), Heterotheca subaxillaris (Camphorweed) and several personal experience statewide and, has access to other experts from several states in the southwest. bibliography

Bass, L.N. 1977. Longevity of seeds of two Dimorphotheca species: Dimorphotheca sinuata
and Dimorphotheca pluvialis. Journal of Seed Technology 2(1):86-87. Bass, L.N., S.J. Toy, R.L. Sayers, and D.C. Clark. 1967. Storage of Dimorphotheca sinuata and Osteospermum ecklonis seed. Proceedings of the Association of Official Seed Beneke, K., M.W. van Rooyen, and G.K. Theron. 1992a. Fruit polymorphism in ephemeral species of Namaqualand: VI. Intermorphic competition among plants cultivated from dimorphic diaspores. South African Journal of Botany 58(6):469-477. Beneke, K., M.W. van Rooyen, and G.K. Theron. 1993c. Fruit polymorphism in ephemeral species of Namaqualand: IV. Growth analyses of plants cultivated from the dimorphic diaspores. Journal of Arid Environments 24:345-360. Beneke, K., M.W. van Rooyen, and G.K. Theron. 1993a. Fruit polymorphism in ephemeral species of Namaqualand: VII. Diaspore production of plants cultivated from dimorphic diaspores. Journal of Arid Environments 25(2):233-247. Beneke, K., M.W. van Rooyen, G.K. Theron, and H.A. van de Venter. 1993b. Fruit polymorphism in ephemeral species of Namaqualand: III. Germination differences between the polymorphic diaspores. Journal of Arid Environments Beneke, K., I. von Teichman, M.W. van Rooyen, and G.K. Theron. 1992b. Fruit polymorphism in ephemeral species of Namaqualand: I. Anatomical differences between polymorphic diaspores of two Dimorphotheca species. South African Burgess, T.L., J.E. Bowers, and R.M. Turner. 1991. Exotic plants at the desert laboratory, Tucson, Arizona. Madroño. 38(2):96-114. de Villiers, A.J., M.W. van Rooyen, G.K. Theron, and H.A. van de Venter. 1994. Germination of three Namaqualand pioneer species, as influenced by salinity, temperature, and light. Seed Science and Technology 22(3):427-433. Earle, W.H. 1973. A new plant for Arizona, Dimorphotheca aurantiaca. Saguaroland GRIN. 2000. Grin Taxonomy. United States Department of Agriculture, Agricultural Research Service, The Germplasm Resources Information Network (GRIN). Website: http://www.ars-grin.gov/npgs/tax/index.html then click on 'simple queries of species data' and search for plant species. Guertin, P. 2001. Observations made during the duration of weed distribution mapping for the USGS Weeds in the West project occurring in the southern Arizona National Park Service management areas. May 1999-June 2001. USGS/BRD, Sonoran Desert Field Station, The University of Arizona, 125 Biological Sciences East, Harris, J.G., and M.W. Harris. 1997. Plant identification terminology: an illustrated glossary. Spring Lake Publishing, Spring Lake, Utah. 197 pp. Hickman, J.C., ed. 1993. The Jepson manual: higher plants of California. University of California Press. Berkeley and Los Angeles, CA. 1400 pp. Holden, M. 1996; unpublished. Exotic plant species list, compiled for southern Arizona parks from park floras and exotic plants lists. National Park Service; Saguaro National Park; Tucson Mountain District; 'soazex~1.xls' database, in MS Excel. Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland: Volume I - Checklist. The biota of North America Program of the North Carolina Botanical Garden. Timber Press, Portland, Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland: Volume II - Thesaurus. The biota of North America Program of the North Carolina Botanical Garden. Timber Press, Portland, Kearney, T.H. and R.H. Peebles. 1960. Arizona flora. University of California Press, Berkeley and Los Angeles, California. 1085 pp. Knowles, R.E., L.A. Glodblatt, G.O. Kohler, S.J. Toy, and J.R. Haun. 1965. Oil seed composition of two species of Dimorphotheca grown at five locations in the United States. Economic Botany 19:262-266. Michigan State University Extension. 1999. Dimorphotheca sinuata -- Cape Marigold. Ornamental Plants plus Version 3.0. Website: http://www.msue.msu.edu/msue/imp/modzz/00000534.html Munz, P.A. 1974. A flora of southern California. University of California Press, Berkeley, Niehaus, T.F., C.L. Ripper, and V. Savage. 1984. The Peterson Field Guide Series. A field guide to southwestern and Texas wildflowers. Sponsored by the National Audubon Society, the National Wildlife Federation, and the Texas Parks and Wildlife Department. Houghton Mifflin Company, Boston, Massachusetts. 449 Rondeau, R., T.R. Van Devender, C.D. Bertelsen, P. Jenkins, R.K. Wilson, M.A. Dimmitt. 1996. Annotated flora and vegetation of the Tucson Mountains, Pima County, Rösch, H., M.W. van Rooyen, and G.K. Theron. 1997. Competitive effect and response of ten Namaqualand ephemeral plant species at two nutrient levels. South African Schulze, B.R. 1965, Climate of South Africa. Part 8. General Survey. W.B. 28. Government Steyn, H.M., N. van Rooyen, M.W. van Rooyen, and G.K. Theron. 1996. The phenology of Namaqualand ephemeral species: the effect of sowing date. Journal of Arid Ueckermann, C., and M.W. van Rooyen. 2000. Insect pollination and seed set in four ephemeral plant species from Namaqualand. South African Journal of Botany United States Department of Agriculture, Natural Resources Conservation Service. 2001. The PLANTS database, Version 3.1 (http://plants.usda.gov/plants/). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. van Rooyen, M.W., N Grobbelaar, G.K. Theron, and N. van Rooyen. 1991. The ephemerals of Namaqualand: effects of photoperiod, temperature, and moisture stress on development and flowering of three species. Journal of Arid Environments van Rooyen, M.W., N Grobbelaar, G.K. Theron, and N. van Rooyen. 1992a. The ephemerals of Namaqualand: effect of germination date on development of three species. van Rooyen, M.W., N Grobbelaar, G.K. Theron, and N. van Rooyen. 1992b. The ephemerals of Namaqualand: effect of germination date on parameters of growth analysis of three species. Journal of Arid Environments 22:117-136. additional sources and websites

Cooperative State Research, Education, and Extension Service
Website: http://www.reeusda.gov/1700/statepartners/usa.htm This website brings you to an interface to connect with Cooperative Extension programs throughout the United States; select the desired state, enter a link, often there is a search option in which information on a plant can be searched for. USDA, Forest Service, Rocky Mountain Research Station, September 2002 has published 'Linking Wilderness Research and Management. Volume 4 - Understanding and Managing Invasive Plants in Wilderness and Other Natural Areas. An Annotated Reading List. General Technical Report RMRS-GTR-79-volume 4 This volume is available on the Web; Website: http://www.fs.fed.us/rm/pubs/rmrs_gtr079_4.pdf (Website: http://www.fs.fed.us/rm/pubs/rmrs_gtr079_4.html provides some information if problems http://ces.asu.edu/collections/vasc_image_library/ImageIndex.jsp then click on appropriate letter, and then scroll down and click on appropriate name

Source: http://sad.hmarts.ru/kat/foto/di/dim_v2.pdf