Cynodon dactylon

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Authors: Dara Newman, Global Invasive Species Team, The Nature Conservancy

Contents


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Taxonomy
Kingdom: Plantae
Phylum: Magnoliophyta
Class: Liliopsida
Order: Cyperales
Family: Poaceae
Genus: Cynodon
Species: dactylon
Scientific Name
Cynodon dactylon
(L.) Pers
Scientific Name Synonym
Panicum dactylon
(L.) Pers
Cynodon aristiglumis
(L.) Pers
Capriola dactylon
(L.) Pers
Common Names

bermudagrass, common bermudagrass, devilgrass

Overview

Appearance
Cynodon dactylon is a perennial grass that occurs on almost all soil types.
Foliage
Leaves are gray-green and 1.5-5.9 in. (4-15 cm) long. The ligule has a ring of white hairs.
Flowers
Flowering occurs in late summer; flowers occur on 1-3 in. (3-7 cm) spikes.
Fruit
This grass spreads by above ground and underground runners.
Ecological Threat
Cynodon dactylon is native to eastern Africa and prefers moist and warm climates with high light. It was introduced into North America in the mid-1800s as a pasture grass. Cynodon dactylon is widely used as a turf grass.

General Description:

Cynodon dactylon is a warm-season, prostrate, perennial grass; it spreads by scaly rhizomes and flat stolons to form a dense resilient turf.

Diagnostic Characteristics:

The distinguishing characteristics of Cynodon dactylon are the conspicuous ring of white hairs of the ligule, the fringe of hairs on the keel of the lemma, and the gray-green appearance of the foliage.

STEWARDSHIP SUMMARY

Cynodon dactylon can be an invasive and competitive weed. The extensive stolon and rhizome system provide a means of rapid expansion. However, this species, which requires high temperatures and high light levels to thrive, grows only in disturbed areas. Although extremely drought tolerant, Bermuda grass tends to grow where water is available. The plant is not frost or shade tolerant and the rhizomes and stolons are susceptible to desiccation. A single treatment or combination of clipping, tilling, shading and herbicide application for several years should result in complete eradication of this weed. The site should be re-vegetated once Bermuda grass is initially controlled in order to prevent the invasion of other weeds or the re-sprouting and establishment of the remaining Bermuda grass rhizomes.

NATURAL HISTORY

Range:

The date of the initial introduction of Bermuda grass into the United States is uncertain, but most likely it occurred in the mid-1800s.[1] By the mid-1900s Bermuda grass had been introduced throughout the southern states and now ranges from California to Florida and occasionally north to Massachusetts and Michigan.[2]

Habitat:

The common name for all the East African rhizomatous species of Cynodon is Bermuda grass.[1][3] Most of these species originated and have remained in southeast Africa. Cynodon dactylon however, has become a "ubiquitous, cosmopolitan weed".[4] The large intra- specific variability in Cynodon dactylon is represented by four varieties which have remained endemic to their original locale, and by two varieties, most notably Cynodon dactylon var. dactylon (from here on referred to by the species name), which have spread to other countries.[1]

Cynodon dactylon grows throughout the warmer regions of both hemispheres. In the United States it occurs at elevations under 6000 feet, primarily in waste places, agricultural fields, and roadsides.[5][6][3] Although widespread, this species "thrives only under extreme disturbance and does not invade natural grasslands or forest vegetation".[4] In areas of low rainfall it commonly grows along irrigation ditches and streambeds.[5][6][3] Bermuda grass, especially the cultivar Coastal, is extremely drought tolerant, however moisture significantly increases its growth rate.[5][7]

Ecology:

Warm temperatures are necessary for the plants to thrive, and long periods of freezing weather or short durations of extremely low temperatures are detrimental to the plants.[8] Average daily temperatures above 24° C are necessary for substantial growth and temperatures of 38° C result in maximum growth rates.[9][3]

In addition to high temperatures, Bermuda grass requires high light intensities. Cynodon dactylon needs direct sunlight in order to grow and dies out with increased levels of shade. This characteristic can be utilized in the control of Bermuda grass.[9]

Cynodon dactylon tolerates a wide range of soil types and conditions. Growth is greater on heavy clay soils than on light sandy soils in dry regions; this may be due to the greater water holding capacity of the clay.[3] Bermuda grass can survive long periods of flooding, but little to no growth occurs without adequate soil aeration.[3] It grows on soils with a wide range of pH values, however alkaline soils are tolerated more than acidic ones. Growth is promoted by the addition of lime to soils with a pH of 5.5.[3] A large amount of available nitrogen is required for maximal above-ground growth; this element is often the limiting factor for Cynodon dactylon. Nitrogen fertilizers are routinely used in order to increase the forage and turf value of Bermuda grass.[5]

The drought and alkali tolerance, and high temperature and sunlight requirements of Bermuda grass explains its success in the Southwest; it is the most common and best performing grass in Arizona.[8] In southern Arizona Cynodon dactylon grows abundantly along sandy washes and near alkaline seeps.[8] A rapidly growing variety, which can grow over hedges 2 m tall, was introduced to Hawaii and Arizona in the early 1900s. A substantial amount of the world's salable seeds of this "giant" Bermuda grass is grown near Yuma, Arizona.[10][4] Large amounts of Bermuda grass, including the giant-type, grow along the edge of Roosevelt dam in Arizona, where it survives submergence under water for part of the year and provides food for cattle during the dry periods.[11]

Shading drastically affects both above- and below-ground growth.[12][13][14][15][16] In Georgia, forage yield is dramatically reduced after the middle of September, with an average June yield of 2907 kg/ha and an average October yield of 295 kg/ha.[15] Daylength and solar radiation, but not rainfall and minimum temperature, were significantly correlated with forage yield; 64%, 43% and 29% of the normal light intensity resulted in a reduced annual dry matter yield of 68%, 42% and 30%, respectively, of unshaded plants.[15] Half the amount of rhizome and root growth occurred in the 64% shade treated-plants than in the control plants.[16] Tall dense trees greatly reduce Bermuda grass growth, and complete canopy cover eventually kills the grass.[12][16] The decrease in growth due to shading is intensified by high temperatures; this may be explained by an increase in respiration rate relative to photosynthetic rate.[13] Increasing the level of nitrogen while maintaining a low light setting results in a further reduction in growth; nitrogen fertilizer increases the retarding effect of low light on shoot, root and rhizome yield, and decreases the amount of reserve carbohydrates while increasing the amount of crude proteins.[16]

Carbohydrate levels in Bermuda grass do not generally follow a consistent pattern.[17] The cultivar and the environmental conditions greatly influence the reserve carbohydrate quantity and quality. In general, total available carbohydrates in the rhizomes increases in the fall, peaking between November and December, decreases in late winter and begins increasing in late spring, reaching a second, but lower, peak in May, and then decreasing in the summer.[18] In Mauritius, the carbohydrate reserves do not decrease in the late winter, instead they increase steadily from fall to spring, and then the pattern fluctuates the rest of the summer depending on the variety.[19] Seasonal rhizome bud germination does not appear to be correlated with the carbohydrate level.[18]

Temperature affects the level of carbohydrates by altering the ratio between the respiration and photosynthetic rate, thus influencing the growth rate.[20] The greatest amount of growth occurred at 30° C/24° C (day/night temperatures) whereas the greatest amount of starch in the stem bases and rhizomes of Coastal Bermuda grass occurred in the 13° C/7° C treated-plants.[20]

Increasing the level of nitrogen results in a decreased amount of reserve carbohydrate.[21][16] Nitrogen fertilizers increase the glucose in leaves by decreasing the amount of sucrose and fructosan in stems, stolons, rhizomes and roots.[22] Nitrogen fertilizers are used to increase the above ground growth of Bermuda grass. When nitrogen is limiting, and the growth conditions unfavorable, fructosans accumulate in the rhizomes.[22] Storage carbohydrate utilization in nitrogen metabolism is thus connected with increased shoot growth.[12]

An increase in nitrogen fertilizer from 0 to 900 pounds per acre results in an increase in height (2.5 inches to 6.5 inches), percent protein, yield (1.6 tons to 11.0 tons of hay), stem length (6.0 to 17.0 inches), internode length and node number, and a decrease in leaf percentage and seed head frequency (5% to 2%).[23] An increase in nitrogen from 0 to 80 kg/ha results in a 5 times greater above-ground biomass.[24]

Bermuda grass is susceptible to desiccation. Long rhizome fragments and dormant stolons require long periods of drying in order to destroy the activity of the buds.[25][26] Air drying of one-node rhizome fragments for seven days resulted in the inhibition of sprouting and a 53% weight loss, however three-node rhizome fragments continued to sprout after seven days of desiccation.[26] Actively growing stolons are more susceptible to desiccation than post-dormant stolons. Greater than 48 hours of drying over an ammonium chloride solution kills actively growing stolons, whereas greater than 96 hours is required to destroy post-dormant fragments.[25] The critical moisture level for stolons is 39% and for the harder to control rhizomes 15%.[25][26] Bermuda grass rhizomes cannot be drowned. Submergence of fragments for eight days in running water or four weeks in stagnant water had no effect on sprouting ability.[26] Thus water is likely to be an efficient means of spreading rhizomes.

COMPETITIVE RELATIONSHIPS:

Plant residues and actively growing plant parts of Cynodon dactylon may pose a direct threat to the growth of neighboring plants.[27][28] Light textured soils mixed for four months with extracts from decaying Bermuda grass plants caused an inhibition of radicle elongation in barley and mustard seedlings.[28] Incubation of test plants for two months with Bermuda grass results in a high degree of inhibition.[27] In addition to the importance of the duration of exposure, is the concentration. The inhibition is proportional to the concentration of plant material. In general, root growth and germination are both affected by decaying residues and actively growing Cynodon dactylon plants.[27] The importance of the allelopathic substances produced by Bermuda grass in the field is unclear. Threats due to completely decayed residues should not be overlooked.

In addition to the allelopathic effects of Cynodon dactylon is the direct competition for space and nutrients by this rapidly growing perennial grass. Bermuda grass's notoriety as a tremendous colonizer comes from the spreading ability of both the rhizomes and stolons.[29] The open growth pattern of Bermuda grass's stolons provides for greater land coverage than seen with species which lack stolons, such as Sorghum halepense; the average monthly area increase in the warm season for Cynodon dactylon and Sorghum halepense is 1.6 m2 and 1.3 m2, respectively.[29] Aerial growth from shoots, tillers and previous season's rhizomes produce an abundance of stolons, which in turn produce more shoots, rhizomes and roots.[30] This growth pattern explains the tremendous spreading capacity of Bermuda grass; the highest monthly area increase was 6 m2 during July and August.[30] However, the average area increase for Cynodon dactylon is only 0.9 m2 per month. This growth rate is far less than other perennial grasses; Cyperus rotundus has a mean area increase of 2.8 m2 per month.[30]

Rhizomes grow in the same configuration as the above-ground growth and are not found growing outside of the sod perimeter. The subterranean dry weight averages 0.6 kg per m3 of soil within a 1 m radius from the center of the plant. Rhizome depth is comparable under the center of the plant and at the edge of the sod. The depth of penetration is restricted by compaction and aeration.[30] With roots extending from stolons and rhizomes, a vast area can be utilized for uptake of water and nutrients.

The competitive ability of Cynodon dactylon depends on the competing plant species and the nutritional level of the soil. Bermuda grass yields were reduced by 40%, 27% and 13% when grown with Johnson grass, Torpedo grass and Cogongrass, respectively for one year.[31] However Bermuda grass had a greater inhibitory effect on the competing plants, with a reduction in the yield of Johnson grass, Torpedo grass and Cogon grass by 55%, 38% and 43%. After two years of competition Johnson grass reduced the yield of Bermuda grass by 99%.[31] Native vegetation recovery, due to the competitive ability of knotgrass (Paspalum sp.), began within one year after cattle were removed from a riparian ecosystem in which Bermuda grass was abundant (Richter pers. comm.).

Cynodon dactylon and Acacia smallii were grown in mixed and mono-culture plots, with and without added fertilizer in order to study the competitive ability and mechanism of the two species.[32] C. dactylon grew 1.5 to 2.4 times larger in mixed cultures than in mono-cultures, with a yield increase of 30% to 50% when grown with Acacia. Cynodon dactylon's competitiveness is thought to stem from its ability to reduce the level of nutrients to below the necessary amount needed by Acacia smallii; this assumption is based on the increase in Cynodon dactylon's growth in the mixed over the mono-culture treatments, its drastic increase in the fertilized, mixed culture plots, and the growth reduction by 70% to 90% of Acacia smallii in the fertilized mixed plots.[32]

Studies on competition in mixed plots of wheat and Cynodon dactylon showed similar intra- and inter-specific competition for nutrients when plants were planted at the same time.[33] Stunted growth of Cynodon dactylon occurred in high wheat density plots. A reduction in dry weight, leaf area and seed output was most likely due to the large size of the wheat plants which caused shading of the Bermuda grass. Low density mono-culture plots of Cynodon dactylon promoted early vegetative spreading growth with delayed reproductive development, whereas in high density plots the period of vegetative growth was shortened and floral development was hastened. In addition, seed production decreased with increased densities of Bermuda grass.[33]

RESPONSE TO MANIPULATIONS AND ABIOTIC FACTORS:

The effects of fire on Cynodon dactylon are variable and dependent on the season and prevailing environmental conditions at the time of burning. Odum et al. (1973)[34] burned a four year old fallow field during the late winter in Georgia. The burn resulted in a drastic reduction in Bermuda grass from 14.7 g/m2 to 0.2 g/m2 as compared to the increase in Johnson grass from 0.2 g/m2 to 27.4 g/m2. However, with the exception of extremely dry conditions or long periods of fire suppression, both of which result in hot fires that may damage the rhizomes, most rhizomatous grasses, such as Cynodon dactylon tend to benefit from fire.[35] Winter burning of Bermuda grass is performed in several southern states in the U.S. in order to hasten spring growth, resulting in increased yield and quality of forage.[36]

The increase in the amount of Cynodon dactylon due to cattle grazing is well documented.[21][17][37][38] Unlike many other plants, intensive grazing on Bermuda grass results in an increase in carbohydrate accumulation in the below-ground structures.[17] This explains the rapid regrowth and establishment that was seen in overgrazed plots in the Serengeti National Park.[37][38] Grazing does not significantly affect growth of rhizomatous and stoloniferous plants that have a prostrate growth form.[21]

Clipping may have a greater affect than grazing on Cynodon dactylon due to the potential for removal of all tillers and shoots.[21] The mowing of Bermuda grass three times a week throughout the growing season had no significant effect on the carbohydrate content or weight of the rhizomes and roots; however, systematic cutting of each individual aerial structure with a scissor resulted in a significant reduction in the reserve carbohydrate level and weight of the below ground structure.[17] Removal of greater than 40% of the shoots reduced root growth and many roots failed to resume growth when severely clipped.[39] Plants with prostrate growth and high, fluctuating levels of reserve carbohydrates, such as Cynodon dactylon, are difficult to control by clipping.[17]

Reproduction:

VEGETATIVE GROWTH:

The principle means of propagation of Cynodon dactylon is through the rhizomes and stolons.[9] These structures are often severed from the plant by burrowing animals and animal hooves; the fragments are then transported by contaminated animals, hay, and machinery, as well as by running water.[9]

The following is a description of the general life-cycle of Cynodon dactylon. In the spring when the temperature begins to increase new stolons elongate and aerial shoots sprout[30]. The characteristic prostrate growth of Bermuda grass lasts for one to several months, early in the season, before flowering culms develop. Most of the lateral growth, produced in concentric circles from the original rhizome, occurs throughout the summer (Horowitz 1972b,).[9] [30]

Single-bud rhizomes were planted and monitored throughout several growing seasons.[19] In the first month a primary shoot and four roots develop from the rhizome. Elongation of the internodes on the shoot is followed by the development of up to twenty buds per node. As many as 12 tillers sprout and three dormant rhizome buds develop from these shoot buds. Horizontal growth commences when the primary shoot and tillers reach 10 cm to 15 cm long, resulting in the formation of stolons. New stolons and roots are continually formed at the nodes of the spreading stolons. The dormant rhizome buds at the basal node of the primary shoot begin to grow at the commencement of the wet warm season. These rhizomes growing deep in the soil provide an over-wintering structure as well as, when they surface, additional above-ground growth.[19]

Rhizomes are the primary over-wintering structure.[9] Cynodon dactylon's success as a weed is thought to be a result of the adaptive rhizome characteristics. The rhizome system is superficial as well as deep, which may account for the ability of this species to infest both arable and waste lands in a variety of conditions. Horowitz (1972a)[30] found approximately 70% of the rhizome weight of two and a half year old plants in the upper 20 cm of the soil and no rhizomes below 40 cm.[30] Other investigators report the existence of rhizomes 1 m deep.[9] Concentric growth, outward from the original rhizome sprout, of the rhizomes corresponds to the circular above-ground growth pattern.[30] A decrease in rhizome dry weight correlates with an increase in distance from the original rhizome. Greater than 60% of the subterranean weight (600 g/m3) occurs within 1 m of the plant center, 30% in the next meter, with the remaining 10% in the 2 to 3 m range.[30] This growth pattern ensures both rapid spreading and strong establishment of the plants.

New rhizomes form only at temperatures greater than 15° C to 20° C; sprouting of rhizome buds is maximal at temperatures between 23° C and 35° C and is inhibited by temperatures below 10° C.[18] No dormancy period is found in rhizomes; sprouting occurs once apical suppression is relieved by fragmentation.[18] Growth of rhizome buds varies depending on depth in soil and age of rhizome. The year-long average sprouting rate for rhizomes in the top 15 cm of soil is 34% and for deeper rhizomes 24%.[18] Young rhizomes sprout much more readily than do older ones. New rhizomes are capable of growing once several distinct internodes have developed. Subterranean growth begins earlier in the spring than aerial growth. The rhizome dry weight increases in April, followed by rhizome elongation.[18] The total rhizome length of single node rhizome fragments planted in July increases from 36 m per plant in December to 95 m the following July, indicating greater rhizome growth in early summer than in late summer.[18]

New stolons are formed at the basal node of shoots which had developed from over-wintering rhizomes.[30] New stolons can grow 75 cm in the first six weeks after sprouting.[30] Within three and a half months eight stolons from the main shoot and seven stolons from secondary shoots develop, resulting in 570 cm of stolon length.[30] The initial stolons move away from the center shoot in straight lines. This is followed by the secondary stolons growing in all directions to form dense circular mats of sod.[30] In two and a half years the average sod area of a single plant is 25 m2, with a maximum growth rate of 2 m2 per month in the summer months.[30] Above ground growth is limited by temperatures below 18° C, and dies at -2° C.[18] The high temperature requirements explain the five times greater increase in sod area for June through November than for December through May.[30]

Roots develop from rhizomes and stolons. Roots produced at the distal end of the stolon are much longer and more abundant than those close to the original stem.[19] Depending on the cultivar, soil texture and nitrogen availability roots can reach 245 cm below the soil surface, however the majority of roots are found in the top 30 cm.[40]

SEXUAL GROWTH:

Asexual reproduction, not sexual reproduction, is responsible for the spread of most Bermuda grass. Most biotypes are infertile, and those that are fertile tend to produce sparse amounts of seeds.[19][9] However, southwestern United States varieties often have a good seed set. The tiny seeds remain viable after passing through livestock and after submergence in water for 50 days.[9] There are approximately 4.4 million seeds in one kilogram.[9] In California, 450 kg of hulled seeds are harvested per hectare in July.[3] All biotypes, regardless of their fertility, produce inflorescences which range in height from 5 cm to 40 cm.[19] Inflorescences develop on the center shoot one and a half months after late spring planting.[30] Inflorescences form during the summer and their production terminates in November; flowers occur throughout the sod at the end of the season, with a maximum of 99 inflorescences per plant. The sod area is proportional to the number of inflorescences with an average of 78 inflorescences per square meter of sod the first year. The second year of growth results in a drastic increase in the number of inflorescences with the maximum of 1125 per plant and an average of 87 inflorescences per square meter of sod.[30]

Impacts:

Spreading of Cynodon dactylon is exacerbated by the continued planting of this turf and forage crop. Bermuda grass is difficult to control once it has been nurtured and has become established.[9][3] The rhizomatous and stoloniferous growth and, depending on the cultivar, the abundance of minuscule seeds, leads to the extensive spreading capabilities of Bermuda grass. However, Bermuda grass is sensitive to shade and frost damage, and only invades disturbed land. Thus, although abundant throughout the world, the threat from the invasion of this plant is limited to warm, sunny, disturbed sites.

Bermuda grass's high pollen production leads to its notoriety as a major contributor to allergies.[8] Forty years ago Bermuda grass was considered the leading cause of hay fever in Arizona.[10]

In addition to competing with native plants for nutrients, Cynodon dactylon presents a direct threat to agricultural crops and possibly to natural vegetation by acting as an alternate host to eleven arthropods, twelve nematodes and numerous viruses.[9][41]

CONDITION

Restoration Potential:

Although Cynodon dactylon is considered the world's weediest grass, eradication appears to be feasible. Bermuda grass poses no problem in undisturbed, cold and shady areas. Thus if land adjacent to invaded areas remains covered by natural vegetation, Bermuda grass will be unable to spread into it.

MANAGEMENT/MONITORING

Management Requirements:

Cynodon dactylon, in disturbed sites, is a competitive and invasive weed. The best management strategy is to remove all plant parts at first sighting. Invasion will be limited by tall plants. Spot herbicide and manual tilling may be adequate in controlling native fields with patchy weed distribution. A more drastic control plan is necessary in sites heavily infested with Bermuda grass. The appropriate manipulation is dependent on the location, humidity, temperature, soil type and precipitation at the specific site. Burning, herbicide application, clipping and shading have all been effective in controlling Bermuda grass under various conditions. In order to prevent the sprouting and establishment of the remaining Bermuda grass rhizomes, native plants and shade material should be installed immediately after the eradication stage.

The best management practice is to avoid the initial invasion of Cynodon dactylon by limiting soil disturbances and maintaining a vegetation cover. Areas where the soil and native plants are kept intact should have little problem from Bermuda grass since it mainly invades disturbed lands (Crosswhite pers. comm., Diamond pers. comm., McWhorter pers. comm., Weigel pers. comm.).

Management strategies depend on the extent of Bermuda grass and the height of the native vegetation.

1a. Sites with established Bermuda grass where restoration projects include re-vegetating with tall-stature plants: A single season of controlling the Bermuda grass following by the transplanting or seeding of native plants will most likely be a sufficient control measure. Periodic spot control or shade mats placed around the young plants may be necessary during the establishment of the native plants. As already stated, the control technique employed is dependent on the site parameters. Herbicide application in conjunction with tilling and desiccation may be the most effective control technique.

1b. Sites with established Bermuda grass where restoration projects include re-vegetating with short-stature plants: A more severe eradication procedure is necessary when the native vegetation will not shade the Bermuda grass. Several repeated tilling and herbicide applications may be required to remove the maximum amount of underground rhizomes and stolons prior to the re-vegetation phase. Many years of spot control may be required until all remaining Bermuda grass is removed or until the ground is covered by the native vegetation.

2a. Uninfested sites with complete canopy cover surrounded by areas containing Bermuda grass: Most likely this situation will not require any active management since Bermuda grass rarely invades undisturbed sites.

2b. Uninfested sites with bare or unshaded spots surrounded by areas containing Bermuda grass: If the site is undisturbed than most likely Bermuda grass will not invade it. However, careful monitoring may show this to not be the case. If appropriate, planting of tall plants between the invaded and non-invaded sites may prevent spreading into the exposed area. Removal of the weed in the adjacent land may be required if invasion occurs frequently.

Several techniques are helpful in controlling Bermuda grass. Depending on the extent of coverage by the weed and on the site parameters a combination of the following manipulations may aid in controlling Bermuda grass: mowing and clipping, tilling and plowing, burning, shading, and chemical control.

MOWING AND CLIPPING:

Removal of the aerial portion of perennial plants may slow the growth by limiting the accumulation of carbohydrates.[42] The temperature, moisture and clipping frequency influence the amount of subsequent growth. Cutting Bermuda grass on hot, dry days has a much greater inhibitory effect than cutting on cool, moist days.[23][43] Weekly clippings at soil level during the moist season reduced yield by 50% whereas clipping during the dry season reduced the yield by 65%.[23] Monthly clipping of Bermuda grass reduced the amount of regrowth in the following year, whereas bi-weekly clipping from spring through winter resulted in the complete inhibition of regrowth the following year.[44] In addition to reducing the regrowth of shoots, the initial clipping inhibited the formation of flowering stalks.[43][44] The removal of aerial portions of the plant as a control measure is only efficient on small scale problem sites due to the labor and time intensity of the necessary frequent repeated clippings.

Eliminating aerial growth reduces the carbohydrate availability in the rhizomes. However understanding the annual carbohydrate cycle is not helpful since rhizome sprouting is not correlated with carbohydrate levels, thus clipping should proceed throughout the entire growing period.[18] The depletion of carbohydrates is related to nitrogen concentrations. Nitrogen fertilizers decrease the amount of carbohydrate reserves; clipping increases the translocation of nitrogen from the soil to the roots and carbohydrate reserves are depleted during the above-ground utilization of nitrogen.[21][24] Care must be taken to remove all aerial growth repeatedly throughout the growing season when clipping and mowing are used as a control measure.

TILLING AND PLOWING:

Hand hoeing is practical only where the concentration of Bermuda grass is low. Shallow cultivation using sharp hoes, shovels, knives or hand pulling will remove the plants and rhizomes from the upper portion of the soil without dividing or pulling up deep rhizomes.[42][45] This technique is impractical in large scale infestations. Repeated plowing throughout the summer growing period will fragment the rhizomes and bring them to the surface; this will aid in the desiccation of the rhizomes and stolons.[3] Small actively growing rhizome and stolon fragments are susceptible to drying within one week.[25][18] Hot, dry weather facilitates desiccation. An alternative to desiccation is freezing. Tilling in the winter will expose the rhizomes to freezing temperatures.[42]

BURNING:

Inconsistent results have been obtained on the effects of burning as a control for Bermuda grass. In general, if conducted at the correct time, burning will slow down the growth of perennial grasses.[42] A late winter fire in Georgia drastically reduced the amount of Bermuda grass.[34] However most burning experiments conducted in swampy areas result in the increase in abundance of Bermuda grass.[35] In wet areas, the rhizomes are protected from the heat of the fire. Due to the variable outcome of burning, this method is not recommended for controlling Cynodon dactylon.

SHADING:

This plant requires high light intensity to thrive. With high levels of shade the plant can no longer grow; thus shading can be used as a control method.[10] Increased amounts of shade results in a decrease in the following: underground carbohydrate level, root weight, rhizome weight and herbage yield.[43] Plants grown under 65% shade resulted in a 68% reduced yield.[15] Results on the survival of Bermuda grass growing under trees indicate that the grass will die when completely shaded by closed canopies[15]. Successful weed control resulted from the use of mats (bought at garden supply shops) which cover the ground and shade the Bermuda grass growing around the base of irrigated trees (Tiller pers. comm.). A thick layer of organic or inorganic mulch may provide adequate shading.

CHEMICAL CONTROL:

Herbicides are helpful in controlling Bermuda grass. However, pre-emergence herbicides are not recommended. These reduce the competition of annual grasses allowing the rhizomes and stolons of Bermuda grass to thrive.[9] An increase in the growth of Bermuda grass is seen in fields where pre-emergence herbicides are used to control annual weeds.[30]

Glyphosate (commercial name -- Roundup®, produced by Monsanto) is mildly toxic and decays rapidly in the soil. This foliar spray, which should not be used in galvanized steel sprayers, is absorbed in the leaves and translocated to growing regions throughout the plant.[46][47] Glyphosate, sprayed from helicopters, trailer sprayers or backpack sprayers, at a concentration of 2% will result in an 85% to 95% control after the first year (Hamilton pers. comm., Heathman pers. comm., Silberman pers. comm., Wildman pers. comm., ).[47]

The best time to spray is when the carbohydrates are being translocated down to the rhizomes at the time of maximum rhizome growth. A downward movement of the herbicides most likely coincides with the spring and fall rhizome growth period.[48] Silberman (pers. comm.) and Brookbank (pers. comm.) recommend the fall spraying of herbicides for maximum effectiveness.

A successful restoration project, restoring cottonwoods and willows to a 40 acre Bermuda grass pasture in California, resulted in maximum control with glyphosate alone (Tiller pers. comm., Silberman pers. comm.). Combinations of herbicides and tilling were less effective. In September while the plants where in full bloom, one week before spraying 2% glyphosate from a boom-sprayer, the field was irrigate to encourage growth. A 95% control was seen after the single application. Trees were planted later, placed on a drip irrigation system and fertilized. Hand weeding and spot herbicide treatment continued for the following year. The greatest regrowth occurred around the irrigated trees. Four feet by four feet shade mats (mulch) were placed around the trees to reduce the sprouting of the remaining rhizomes.

Management Programs:

The following people are involved in either actively managing or planning the management of Cynodon dactylon:

  • Val Little, Preserve Manager, Hassayampa River Preserve, The Nature Conservancy, Box 1162, Wickenburg, AZ 85358; (602) 684- 2772.
  • Oren Pollack, Stewardship Ecologist, California Regional Office, The Nature Conservancy, 785 Market St., 3rd Floor, San Francisco, CA 94103; (415) 777-0487.
  • Ron Tiller, Preserve Manager, Kern Preserve, The Nature Conservancy, P.O. Box 1662, Weldon, CA 93283; (619) 378-2531.

Monitoring Requirements:

Monitoring the size of the area of land infested by Bermuda grass would be beneficial in determining the optimal control technique. Since Bermuda grass is low growing and not always readily apparent, some type of marking system should be employed in order to expedite the yearly measurements. Knowledge of the extent of the underground rhizome and root system is important for manipulating the entire infested area; no additional measurements are required since the underground growth pattern parallels the aerial development.

Yearly summer monitoring of Bermuda grass should determine whether the aerial extent is diminishing with the employed control measure. Tagging the edge of each cluster yearly may help in visually assessing the expansion or reduction in the infested area and in rapidly locating the problem site.

Both the sod area and maximum extension are useful measurements. Changes in sod area can be determined by comparing the area of the annual concentric circle growth size drawn on a contour map, plotted by making a grid with poles spaced 1 m apart and then subtracting the size of the bare areas.[30] A random sampling of segments of the infested field may provide sufficient information for large scale problems. Maximum extension is measured by determining the distance from the center of the sod to the tip of the furthest stolon. Measuring the number of inflorescences produced each summer is helpful with fertile varieties.[30] Germination tests will determine the fertility status of the variety in question.

Monitoring Programs:

Presently no formal monitoring programs of Bermuda grass are known. The following people "eyeball" the distribution of the grass:

  • Val Little, Preserve Manager, Hassayampa River Preserve, The Nature Conservancy, Box 1162, Wickenburg, AZ 85358; (602) 684- 2772.
  • Ron Tiller, Preserve Manager, Kern Reserve, The Nature Conservancy, P.O. Box 1662, Weldon, CA 93282; (619) 378-2531.
  • Jeff Weigel, Director of Stewardship, The Nature Conservancy, P.O. Box 1440, San Antonio, TX 78295-1440; (512) 224-8774.

RESEARCH

Management Research Programs:

No specific research on Bermuda grass control in natural plant communities is being conducted presently. However, ongoing research on the eradication of perennial weedy grasses with the use of newly synthesized herbicides in the agricultural milieu takes place in most of the southern land-grant universities (Hamilton pers. comm.).

Management Research Needs:

An extensive amount of information on Cynodon dactylon is available. The genetics, life-cycle, environmental requirements, phenology, beneficial and deleterious characteristics, and control of Bermuda grass are all well documented. However, the information pertaining to controlling this species pertains to agricultural crop fields and not to the natural environment. Most of these techniques are not economically or practically feasible in a non-agricultural setting.

Information on controlling Bermuda grass in a natural setting is needed. Two major stages, not necessarily temporally separated, are essential for restoring the native flora: eradication of the weed and encouragement of native plants, preferably large, if appropriate. The extensive rhizome system essentially prevents the complete removal of Bermuda grass, thus once controlled, periodic manipulation of the weed is necessary. The early establishment of native plants which can shade the Bermuda grass is important to eradication, maintenance and re-vegetation.

Information on both optimal manipulation and native competitor establishment must be specific for the problem site. The temperature, precipitation, humidity and elevation will determine the optimal control technique.

Experimental plots should be employed for long term studies of various manipulation techniques. Controlled burning at various times of the year and assorted repetition cycles from single burns to yearly repeated burns should be analyzed (Cox pers. comm.). Various schedules of mowing, grazing, tilling, desiccating and herbicide applications should be studied at different locations. The percent coverage, timing of shading and types of shading material, such as shade cloth, shade mats, trees and other plants, should be studied in order to maximize the shade sensitivity of the species. The effects on carbohydrate reserves of nitrogen amendments to shaded plants should be analyzed. The interaction of several manipulation techniques should also be examined.

Rapid recovery of native vegetation, once most of the Bermuda grass has been removed, is essential in order to prevent invasion by other weeds or re-sprouting and establishment of the remaining Bermuda grass rhizomes. Studies to determine the optimal native species to be used and re-vegetation schedule to be followed must be conducted.[49] If appropriate, the establishment of large native plants will provide both a shading device and re- vegetation material.

Resources

INFORMATION SOURCES

Bibliography

  1. Harlan, J. 1970. Cynodon species and their value for grazing and hay. Herbage Abstract 40:233-238. 1.0 1.1 1.2
  2. Hitchcock, A. S. and A. Chase. 1950. Manual of the grasses of the United States, second edition. USDA miscellaneous Publication No. 200. United States Government Printing Office, Washington. 1051 pp.
  3. Burton, G., and W. Hanna. 1985. Burmudagrass. In M. Heath, R. Barnes, and D. Metcalfe ed. Forages the science of grassland agriculture. Iowa State University Press, Ames, Iowa. 643 pp. 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9
  4. Harlan, J., and J. de Wet. 1969. Sources of variation in Cynodon dactylon. Crop Science 9:774-778. 4.0 4.1 4.2
  5. Humphrey, R. 1977. Arizona range grasses; their description, forage value, and management. University of Arizona Press, Tucson, Arizona. 159pp. 5.0 5.1 5.2 5.3
  6. Crampton, B. 1974. Grasses in California. University of California Press, Berkeley, California. 178 pp. 6.0 6.1
  7. Burton, G., J. Butler, and R. Hellwig. 1987. Effect of supplemental irrigation on the yield of coastal Bermudagrass in the southeastern United States. Agronomy Journal 79:423-424.
  8. Gould, F. 1951. Grasses of southwestern United States. University of Arizona, Tucson, Arizona. 343 pp. 8.0 8.1 8.2 8.3
  9. Holm, L. G., P. Donald, J. V. Pancho, and J. P. Herberger. 1977. The World's Worst Weeds: Distribution and Biology. The University Press of Hawaii, Honolulu, Hawaii. 609 pp. 9.00 9.01 9.02 9.03 9.04 9.05 9.06 9.07 9.08 9.09 9.10 9.11 9.12
  10. Kearney, T.H., and R.H. Peebles. 1951 (with supplement in 1960). Arizona Flora. Univ. California Press, Berkeley. 1085 pp. 10.0 10.1 10.2
  11. Munz, P.A. (in collaboration with D.D. Keck). 1959. A California flora. Univ. California Press, Berkeley. 1681 pp + supplement.
  12. Schmidt, R., and R. Blaser. 1969. Effect of temperature, light, and nitrogen on growth and metabolism of 'tifgreen' Bermudagrass. Crop Science 3:5-9. 12.0 12.1 12.2
  13. McBee, G., and E. Holt. 1966. Shade tolerance studies on Bermudagrass and other turfgrasses. Agronomy Journal 58:523-525. 13.0 13.1
  14. Hart, R., R. Hughes, C. Lewis, and W. Monson. 1970. Effect of nitrogen and shading on yield and quality of grasses grown under young slash pines. Agronomy Journal 62:285-287.
  15. Burton, G., J. Hook, J. Butler, and R. Hellwig. 1988. Effect of temperature, daylength, and solar radiation on production of coastal Bermudagrass. Agronomy Journal 80:557-560. 15.0 15.1 15.2 15.3 15.4
  16. Burton, G., J. Jackson, and F. Knox. 1959. The influence of light reduction upon the production persistence and chemical composition of coastal Bermudagrass, Cynodon dactylon. Agronomy Journal 51: 537-542. 16.0 16.1 16.2 16.3 16.4
  17. Weinmann, H. 1961. Total available carbohydrates in grasses and legumes. Herbage Abstracts 31:255-260. 17.0 17.1 17.2 17.3 17.4
  18. Horowitz, M. 1972b. Development of Cynodon dactylon. Weed Research 12:207-220. 18.0 18.1 18.2 18.3 18.4 18.5 18.6 18.7 18.8 18.9
  19. Rochecouste, E. 1962a. Studies on the biotypes of Cynodon dactylon; botanical investigations. Weed Research 2:1-23. 19.0 19.1 19.2 19.3 19.4 19.5
  20. McKell, C. M., B. B. Youngner, F. J. Nudge and J. J. Chatterton. 1969. Carbohydrate accumulation of coastal Bermuda grass and Kentucky bluegrass in relation to temperature regimes. Crop Sci. 9:534-537. 20.0 20.1
  21. White, L. 1973. Carbohydrate reserves of grasses: a review. Journal of Range Management 26(1): 13-18. 21.0 21.1 21.2 21.3 21.4
  22. Adegbola, A. and C. McKell. 1966. Effect of nitrogen fertilization on the carbohydrate content of coastal Bermudagrass (Cynodon dactylon). Agronomy Journal 58: 60-64. 22.0 22.1
  23. Prine, G., and G. Burton. 1956. The effect of nitrogen rate and clipping frequency upon the yield, protein content, and certain morphological characteristics of coastal Bermudagrass (Cynodon dactylon). Agronomy Journal 48:296-301. 23.0 23.1 23.2
  24. Skousen, J., C. Call, and R. Weaver. 1989. Recovery of N15-labelled fertilizer by coastal Bermudagrass in lignite minesoil. Plant and Soil 114:39-43. 24.0 24.1
  25. Webb, B. 1959. Comparison of water loss and survival of coastal Bermudagrass stolons harvested at two stages of growth. Agronomy Journal 51:367-368. 25.0 25.1 25.2 25.3
  26. Horowitz, M. 1972d. Effects of desiccation and submergence on the viability of rhizome fragments of Bermudagrass and Johnson grass and tubers of Nutsedge. Israel Journal of Agricultural Research 22:215-220. 26.0 26.1 26.2 26.3
  27. Horowitz, M., and T. Friedman. 1971. Biological activity of subterranean residues of Cynodon dactylon L., Sorghum halepense L. and Cyperus rotundus L. Weed Research 11:88-93. 27.0 27.1 27.2
  28. Friedman, T., and M. Horowitz. 1970. Phytotoxicity of subterranean residues of three perennial weeds. Weed Research 10: 382-385. 28.0 28.1
  29. Horowitz, M. 1973. Spatial growth of Sorghum halepense. Weed Research 13:200-208. 29.0 29.1
  30. Horowitz, M. 1972a. Spatial growth of Cynodon dactylon. Weed Research 12: 373-383. 30.00 30.01 30.02 30.03 30.04 30.05 30.06 30.07 30.08 30.09 30.10 30.11 30.12 30.13 30.14 30.15 30.16 30.17 30.18 30.19 30.20
  31. Wilcut, J., B. Truelove, D. Davis, and J. Williams. 1988. Temperature factors limiting the spread of Cogongrass (Imperata cylindrica) and Torpedograss (Panicum repens). Weed Science 36:49-55. 31.0 31.1
  32. Cohn, E., O. VanAuken, and J. Bush. 1989. Competitive interactions between Cynodon dactylon and Acacia smallii seedlings at different nutrient levels. American Midl. Naturalist 121:265-272. 32.0 32.1
  33. Ramakrishnan, P., and S. Kumar. 1971. Productivity and plasticity of wheat and Cynodon dactylon in pure and mixed stands. Journal of Applied Ecology 8:85-98. 33.0 33.1
  34. Odum, E., S. Pomeroy, J. Dickinson, and K. Hutcheson. 1973. The effects of late winter litter burn on the composition, productivity, and diversity of a 4-year old fallow-field in Georgia. Tall Timbers Fire Ecology Conference 13:399-419. 34.0 34.1
  35. Rensburg, H. 1970. Fire: its effect on grasslands, including swamps-southern, central and eastern Africa. Tall Timbers Fire Ecology Conference 11:175-199. 35.0 35.1
  36. Hardison, J. 1974. Fire and disease. Tall Timbers Fire Ecology Conference 15:223-233.
  37. Belsky, A. J. 1986a. Revegetation of artificial disturbances in grasslands of the Serengeti National Park, Tanzania; colonization of grazed and ungrazed plots. Journal of Ecology 74:419-437. 37.0 37.1
  38. Belsky, A. J. 1986b. Revegetation of artificial disturbances in grasslands of the Serengeti National Park, Tanzania; five years of successional change. Journal of Ecology 74:937-951. 38.0 38.1
  39. Risser, P., E. Birney, H. Blocker, W. May, W. Parton, and J. Wiens. 1981. The true prairie ecosystem. Huchinson Ross Publishing Co., Stroudsburg, Pennsylvania. 557 pp.
  40. Burton, G., E. DeVane, and R. Carter. 1954. Root penetration, distribution, and activity in southern grasses measured by yields, drought symptoms, and P32 uptake. Agronomy Journal 51:537-542.
  41. Akobundu, I. 1987. Weed science in the tropics; principles and practices. John Wiley and Sons Ltd., NY. 522 pp.
  42. Lorenzi, H., and L. Jeffery. 1987. Weeds of the U.S. and their control. Van Nostrand Reinhold Co., New York. 355 pp. 42.0 42.1 42.2 42.3
  43. Jameson, D. 1963. Responses of individual plants to harvesting. Botanical Review 29(4):532-594. 43.0 43.1 43.2
  44. Horowitz, M. 1972c. Effects of frequent clipping on three perennial weeds, Cynodon dactylon, Sorghum halepense, and Cyperus rotundus. Experimental Agriculture 8:225-234. 44.0 44.1
  45. Heathman, S., K. Hamilton, and J. Chernicky. 1986. Control weeds in urban areas. Cooperative Extension Service 8653. University of Arizona, Tucson, Arizona. 4 pp.
  46. Ross, M. 1986. Johnsongrass- two decades of progress in control. Crops and Soils 39: 12-14.
  47. Kelly, J. 1983. How to convert a Bermudagrass lawn to a gravel or desert landscape. Cooperative Extension Service. University of Arizona, Tucson, Arizona. p.1. 47.0 47.1
  48. Rochecouste, E. 1962b. Studies on the biotypes of Cynodon dactylon; growth response to trichloroacetic and 2,2-dichloropropionic acids. Weed Research 2:136-145.
  49. Newman, D. 1989. Element Stewardship Abstract- Sorghum halepense. The Nature Conservancy, Tucson, Arizona. September 11.

Additional References

  • Sampson, A., A. Chase, and D. Hedrick. 1951. California grasslands and range forage grasses. California Agricultural Experiment Station Bulletin 724:3-130.

Original Document

Element Stewardship Abstract; Dara Newman, 2001.


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