ELEMENT STEWARDSHIP ABSTRACT for

Anthoxanthum odoratum

Sweet Vernal Grass

To the User:

Element Stewardship Abstracts (ESAs) are prepared to provide The Nature Conservancy's Stewardship staff and other land managers with current management-related information on those species and communities that are most important to protect, or most important to control. The abstracts organize and summarize data from numerous sources including literature and researchers and managers actively working with the species or community.

We hope, by providing this abstract free of charge, to encourage users to contribute their information to the abstract. This sharing of information will benefit all land managers by ensuring the availability of an abstract that contains up-to-date information on management techniques and knowledgeable contacts. Contributors of information will be acknowledged within the abstract and receive updated editions. To contribute information, contact the editor whose address is listed at the end of the document.

For ease of update and retrievability, the abstracts are stored on computer at the national office of The Nature Conservancy. This abstract is a compilation of available information and is not an endorsement of particular practices or products.

Please do not remove this cover statement from the attached abstract.

Authors of this Abstract: Don Pitcher, Mary J Russo © THE NATURE CONSERVANCY 1815 North Lynn Street, Arlington, Virginia 22209 (703) 841 5300 The Nature Conservancy Element Stewardship Abstract For Anthoxanthum odoratum

I. IDENTIFIERS

Common Name: SWEET VERNAL GRASS Global Rank: G?

General Description: Anthoxanthum odoratum is a sweet-smelling perennial grass with flat blades and spikelike panicles.

The name Anthoxanthum is derived from the Greek, ANTHOS meaning flower and XANTHOS meaning yellow, alluding to the yellow inflorescence.

Diagnostic Characteristics: Anthoxanthum odoratum can be distinguished from other grass species by its perennial habit and sweet, vanilla scent, especially when dry.

II. STEWARDSHIP SUMMARY

Anthoxanthum odoratum is a sweet-smelling perennial grass of Eurasian origin that has escaped from cultivation in North America. It occurs primarily on poor soils but is not considered a major weed pest. It reproduces by seeds and can be highly competitive with other grasses, particularly during the spring. Sweet vernal grass also shows a remarkable ability to genetically adapt to different environmental conditions. It can be controlled by hand pulling individual plants, hand hoeing, mowing, or by using the herbicide Dalapon. Monitoring needs to be continued after the initial removal of Anthoxanthum odoratum plants because of their potentially long seed dormancy in the soil.

III. NATURAL HISTORY

Range: It ranges from northern Florida to southern Canada along the East Coast and west to the Mississippi River flood plain. It also occurs from northern California to Vancouver Island, Canada (Hitchcock 1971, Grant and Antonovics 1978). It was introduced to North America from Europe in the late 1700s as a meadow grass and has since escaped cultivation (Munz and Keck 1973, Grant and Antonovics 1978).

Habitat: Sweet vernal grass is a native of Eurasia (Hitchcock 1971, Munz and Keck 1973). It occurs on poorer soils in dry fields, lawns, meadows, and waste places (Muenscher 1955).

Ecology: GROWTH AND COMPETITION When grown with other common pasture grasses, Anthoxanthum odoratum is highly competitive. In a North Carolina study, it generally dominated mixtures that included Plantago lanceolata, Poa pratensis, Salvia lyrata, and Rumex acetosella (Fowler 1982).

In a study of competition with Lolium perenne, Dactylis glomerata, and Holcus lanatus (Remison and Snaydon 1978), sweet vernal grass proved highly competitive in the early spring due to its rapid growth and early flowering. Later in the summer it declines in aggressiveness. Among these species, sweet vernal grass is the most highly competitive for potassium (Remison and Snaydon 1978). A Dutch study revealed similar results (Berendse 1983). Even at low densities, competition between Anthoxanthum odoratum and Plantago lanceolata is very strong. Anthoxanthum's competitive ability is highest in early spring.

In competition experiments (Remison 1978) with Holcus lanatus and Dactylis glomerata, Anthoxanthum odoratum had the lowest yield, especially at low densities. It proved most competitive at intermediate densities. Again, however, sweet vernal grass was most competitive early in the growing season, due to its rapid spring growth.

Anthoxanthum odoratum roots are quite shallow, absorbing nutrients mainly from the upper 10 cm of soil (Berendse 1982). It commonly occurs on soils that are low in phosphorus (Roberts 1982). On fertilized soils, sweet vernal grass develops a dense surface root mat (Remison and Snaydon 1978). Although Anthoxanthum tillers profusely, its poor root growth reduces its competitive ability (Remison 1978). The ratio of roots to shoots is therefore quite low at 0.(55) (Remison and Snaydon 1978).

In mixed species plots, intraspecific competition is more important than interspecific competition (Berendse 1983). Like many other grasses, sweet vernal grass contains allelopathic chemicals that suppress the growth of other plant species (Scott 1975). It is "allo-inhibited," growing better with exudate from its own and similar species in permanent grasslands (Newman and Rovira 1975).

The decomposing roots of Anthoxanthum odoratum have an effect opposite that of the allelopathic leachate. Old roots appear to enhance the growth of other grass species, while decreasing the growth of new Anthoxanthum plants. Phosphorus content in the roots averaged 0.155%, a relatively high level. This is probably one of the factors leading to the increased growth of other species on sites with decomposing sweet vernal grass roots (Newberry 1979).

Anthoxanthum odoratum plants average 3.5 leaves per shoot. The individual leaves are unusually short-lived, with a mean survival of only 19 days. Only a few leaves survive for three months. This rapid turnover of leaves is probably a reflection of the plant's rapid growth rate (Sydes 1984).

POPULATION DYNAMICS The biology of Anthoxanthum odoratum populations is complex and depends upon the particular habitat. Different character complexes are favored by different habitat types leading to substantial phenetic differences between ecologically marginal and central populations. Ecologically marginal populations have higher turnover rates than central ones. Individual life expectancy is shorter in marginal populations, while central populations contain a skewed age distribution in favor of older individuals, making them potentially less stable (Grant and Antonovics 1978).

In response to contrasting environments, sweet vernal grass can rapidly differentiate, even without geographic barriers. In one study (Kiang 1982), significant differences in morphology, flowering date, and pollen fertility developed in less than 40 years. Such differences among populations were genetic adaptations to contrasting soil environments. Similar results are reported by Snaydon and Davies (1972, 1976) for populations at distances of less than 30 m apart.

When environmental boundaries are distinct, Anthoxanthum odoratum populations can exhibit sharp differences. Plants growing only 0.1 m apart across a sharp environmental boundary that had existed for less than 120 years showed significant differences in height, yield, flowering date, and morphology (Snaydon and Davies 1976).

Reproduction: Anthoxanthum odoratum reproduces from seeds, flowering early in the spring and forming distinct, identifiable clumps that do not spread vegetatively (Grant and Antonovics 1978). The plants are generally self-incompatible (Antonovics 1972). Plants in North Carolina were reported to produce from 58 to 1,257 seeds per plant (Grant and Antonovics 1978).

SEED DISPERSAL AND GERMINATION Seeds with awns are able to move further along the soil surface than awnless seeds. Anthoxanthum odoratum seeds have awns that wind and unwind with changes in atmospheric humidity. The movement of seeds by this mechanism is not great, averaging 1.6 cm on dry soil and 1.3 cm on wet soil over a 5-day period, but it may give the seeds a slight advantage in reaching suitable germination sites (Schonfeld and Chancellor 1983).

Harris (1961) found some dormancy in Anthoxanthum odoratum seeds. Most germination began the 4th week after sowing and ceased by the 8th week, but a few seeds germinated more than a year later. In experimental studies in England (Williams 1983a), A. odoratum seeds stored for 48 weeks in dry storage resulted in 86% germination. There is thus indirect evidence of seed dormancy, which may increase a seed's chance of being incorporated into the soil while still viable. For seeds sown on moist soil, however, emergence occurred rapidly (within three months). Those seeds that did not emerge within 10 months were not viable. Emergence was highest for seeds sown at a depth of 1 cm (75% viable) and decreased to 52% for seeds sown at a depth of 5 cm.

In another British study (Williams 1983b), Anthoxanthum odoratum was compared with eight other grassland species. It showed a greater ability to germinate under constant conditions than when conditions were varied. When A. odoratum seeds were stored for 28 weeks, a much higher percentage of the seeds germinated when stored at constant temperatures.

IV. CONDITION

Threats: Anthoxanthum odoratum occurs on Nature Conservancy grassland preserves in Oregon from west of the Cascade Mountains to the coast. At present, it is not a significant threat to California preserves, but where present, as at the Northern California Coast Range Preserve, its elimination is the ultimate goal.

Restoration Potential: With the right combination of control measures, it should be possible to eliminate sweet vernal grass from selected areas. Annual monitoring and removal must continue until soil seedbanks are exhausted.

V. MANAGEMENT/MONITORING

Management Requirements: Control of Anthoxanthum odoratum requires active management once it becomes established in an area. Control through prescribed burning is being attempted on Oregon preserves.

MECHANICAL CONTROL The physical removal of sweet vernal grass by hand pulling or hoeing is an effective method of control for small patches. Control can also be obtained by mowing early in the season and before the seeds have matured. Badly infested fields should be plowed and seeded with a cover crop (Muenscher 1955).

BIOLOGICAL CONTROL According to Charles Turner of the USDA Biocontrol Lab in Albany, California (pers. comm. 1985), grass species such as Anthoxanthum odoratum present a major problem for biocontrol techniques. The lack of known host-specific insect or disease pest controls means that more experimenting must be done. Such experiments could damage ecologically or agriculturally important grass species. In addition, sweet vernal grass is not considered an undesirable plant everywhere. Keeping insects or pathogens within property boundaries is impossible.

CONTROL BY GRAZING Grazing is not known to be a useful control method for Anthoxanthum odoratum.

CHEMICAL CONTROL A variety of herbicides are effective on Anthoxanthum, but they will also kill other, more desirable grasses. Dalapon (Dowpon) provides excellent control of Anthoxanthum odoratum (Fryer and Makepeace 1978). Dr. Jim McHenry (pers. comm. 1985), of the University of California, Davis, recommends it to control sweet vernal grass on Nature Conservancy lands. It will kill both annual and perennial grasses but is not as effective on broadleaf plants. Dalapon should be applied in the spring when there is maximum top growth, prior to the appearance of the seed head. Apply the herbicide at the rate of 10 lbs/100 gal of water along with one quart of surfactant. Surfactants lower the surface tension of the spray and increase the herbicide's effectiveness. Dalapon is cleared for use on right-of-ways and grazing lands (if spot treated). Dalapon has low toxicity to animals and fish and breaks down rapidly and completely in soils (Beste 1983).

Herbicides can be applied uniformly over an area for large infestations or by spot spraying individuals plants. Due to the nonselective nature of Dalapon, spot spraying is recommended for most purposes. Dr. McHenry recommends using a flat-fan nozzle (Spraying Systems Co. #8003 or #8004 nozzle tip) rather than the cone nozzles available on most garden sprayers. Cone sprayers produce greater atomization of the chemicals and increase the chance of unwanted drift. Spraying should be done to dry plants on calm days since dew or rain will dilute the herbicide, reducing its effectiveness. When spraying large areas, a horizontal boom (6-8 feet long) made from aluminum tubing will improve herbicide coverage.

Management Programs: Annual monitoring following prescribed burning continues at Cascase Head and Willow Creek preserves in Oregon.

Monitoring Requirements: Monitoring is needed to determine the effectiveness of any sweet vernal grass control measures. Methods will vary depending upon the degree of the problem on a given preserve.

A. odoratum is being monitored on all Oregon preserves where it occurs using nested frequency methods and, in some cases, with density counts (Macdonald pers. comm. 1988). It is not being monitored in California.

Monitoring Programs: For more information on monitoring efforts in Oregon contact: Cathy Macdonald, Land Steward The Nature Conservancy Oregon Field Office 1205 NW 25th Avenue Portland, OR 97210 (503) 228-9561

VI. RESEARCH

Management Research Programs: Research burns were conducted in 1986 and 1987 at Willow Creek Preserve, OR. Additional research burns were done at Cascade Head Preserve, OR, in 1988. It is still too early to evaluate the results of these burns on the control of sweet vernal grass (Macdonald pers. comm. 1988). For further information contact: Cathy Macdonald, Land Steward The Nature Conservancy Oregon Field Office 1205 NW 25th Avenue Portland, OR 97210 (503) 228-9561

Management Research Needs: Very little research has been done on controlling sweet vernal grass. Much research needs to be done in the areas of biological control, prescribed burning, and mechanical removal techniques.

VII. ADDITIONAL TOPICS

VIII. INFORMATION SOURCES

Bibliography:

Abrams, L. 1940. Illustrated flora of the Pacific states: Washington, Oregon, and California. Vol. I. Ophioglossaceae to Aristolochiaceae. Stanford Univ. Press, Stanford, California. 538 pp.

Antonovics, J. 1972. Population dynamics of the grass Anthoxanthum odoratum on a zinc mine. J. Ecology. 60:351-365.

Antonovics, J. 1984. Genetic variation within populations. Pp. 229-241 in R. Dirzo and J. Sarukhan (ed.), Perspectives of plant population ecology. Sinauer Associates, Inc., Sunderland, MA.

Berendse, F. 1982. Competition between plant populations with different rooting depths. III. Field experiments. Oecologia 53:50-55.

Berendse, F. 1983. Interspecific competition and niche differentiation between Plantango lanceolata and Anthoxanthum odoratum in a natural hay field. J. Ecology 71:379-390.

Beste, C. E. 1983. Herbicide handbook. Weed Science Society of America, Herbicide Handbook Committee. Champagne, IL.

Dwire, K. A. 1983. Dispersal and colonization of the invading perennial grass, Anthoxanthum odoratum, in annual patches in a California coastal grassland. MS thesis. Univ. of California, Davis.

Fowler, N. 1982. Competition and coexistence in a North Carolina Grassland. III. Mixtures of component species. J. Ecology 70:77-92.

Fryer, J. D. and R. J. Makepeace. 1978. Weed control handbook. Vol. II. Recommendations. Blackwell Scientific Publications, Oxford, England. Grant, Michael C., and Janis Antonovics. 1977. Biology of ecologically marginal populations of Anthoxanthum odoratum: I. phenetics and dynamics. Evolution. 32(4):822- 838.

Harris, G. S. 1961. The periodicity of germination in some grass species. New Zealand Journal of Agricultural Research 4:253-260.

Hitchcock, A.S. 1951. Manual of the grasses of the United States. 2nd edition revised by Agnes Chase, 1971. 2 vols. Dover Publications, Incorporated, New York.

Jepson, W.L. 1925. A manual of the flowering plants of California. Independent Pressroom: San Francisco, CA.

Kinag, Y. t. 1982. Local differentiation of Anthoxanthum odoratum populations on roadsides. American Midlands Naturalist 107:340-350.

Macdonald, C. 1988. Oregon Land Steward, The Nature Conservancy. Memorandum to M. J. Russo, The Nature Conservancy, Western Regional Office. September 11, 1988.

McHenry J. 1985. University of California, Davis, CA. Personal communication to Don Pitcher.

Muenscher, W. C. 1955. Weeds. The MacMillan Co., New York.

Munz, P.A., and D.D. Keck. 1973. A California flora and supplement. Univ. California Press, Berkeley, CA.

Newberry, D. M. 1979. The effects of decomposing roots on the growth of grassland plants. J. Applied Ecology 16:613-622.

Newman, E. I. and A. d. Rovira. 1975. Allelopathy among some British grassland plants. J. of Ecology 63:727-737.

Remion, s. U. 1978. The effects of decomposing roots on the growth of grassland plants. J. Applied Ecology 16:613-622.

Remison, S. U. and R. W. Snaydon. 1978. Yield, seasonal changes in root competitive ability and competition for nutrients among grass species. J. Agricultural Science, Cambridge 90:115-124.

Roberts, H. A., ed. 1982. Weed control handbook: principles. Blackwell Scientific Publications, Oxford, England.

Schonfeld, M. A. and R. J. Chacellor. 1983. Factors influencing seed movement and dormancy in grass seeds. Grass and Forage Science 38:234-250. Scott, D. 1975. Allelopathic interactions of resident tussock grassland species on germination of oversown seed. New Zealand Journal of Experimental Agriculture 3:135- 141.

Snaydon, R. W. 1970. Rapid population differentiation in a mosaic environment. The response of Anthoxanthum odoratum to soils. Evolution 24:257-269.

Snaydon, R. W. and M. S. Davies. 1972. Rapid population differentiation in a mosaic environment. II. Morphological variation in Anthoxanthum odoratum. Evolution 26:390- 405.

Snaydon, R.W. and M. S. Davies. 1976. Rapid population differentiation in a mosaic environment. IV. Populations of Anthoxanthum odoratum at sharp boundaries. Heredity 37:9-25.

Sydes, C. L. 1984. A comparative study of leaf demography in limestone grassland. J. Ecology 72:331-345.

Tuner, C. 1985. USDA Biocontrol Lab in Albany, California. Personal communication to Don Pitcher.

Williams, E. D. 1983. Effects of temperature, light, nitrate and pre-chilling on seed germination of grassland plants. Annals of Applied Biology 103:161-172.

Williams, E.D. 1983. Germinability and enforced dormancy in seeds of species of indigenous grassland. Annals of Applied Biology 102:557-566.

IX. DOCUMENT PREPARATION & MAINTENANCE

Edition Date: 88-11-04 Contributing Author(s): Don Pitcher, Mary J Russo (Revision), CAFO