Cynodon dactylon
(Bermuda grass)

Pictures

Picture	Title	Caption	Copyright
Title leaves Caption Cynodon dactylon (Bermuda grass); leaves. Puu Moaulanui, Kahoolawe, February, 2008 Copyright ©Forest Starr & Kim Starr - CC BY 4.0	leaves	Cynodon dactylon (Bermuda grass); leaves. Puu Moaulanui, Kahoolawe, February, 2008	©Forest Starr & Kim Starr - CC BY 4.0
Title Shoot and inflorescence Caption Copyright ©Chris Parker/Bristol, UK	Shoot and inflorescence		©Chris Parker/Bristol, UK
Title Inflorescence and leaves - line drawing Caption Inflorescence supported on a culm up to 25 cm high, consisting of a single whorl of 3-7 narrow racemes, each 3-8 cm long. Spikelets 2-2.5 mm long, in two rows, closely appressed to the rachis. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: CIBA-GEIGY. Copyright NOVARTIS	Inflorescence and leaves - line drawing	Inflorescence supported on a culm up to 25 cm high, consisting of a single whorl of 3-7 narrow racemes, each 3-8 cm long. Spikelets 2-2.5 mm long, in two rows, closely appressed to the rachis. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: CIBA-GEIGY.	NOVARTIS
Title Spikelet - line drawing Caption Glumes one-nerved, lower almost as long as the spikelet, the upper, half to three quarters as long. Lemma silky pubescent on the keel; palea glabrous. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: Ciba-Geigy. Copyright ©Chris Parker/Bristol, UK	Spikelet - line drawing	Glumes one-nerved, lower almost as long as the spikelet, the upper, half to three quarters as long. Lemma silky pubescent on the keel; palea glabrous. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: Ciba-Geigy.	©Chris Parker/Bristol, UK
Title Blade, ligule and sheath - line drawing Caption Ligule very short, 0.2-0.3 mm, with a conspicuous fringe of white hairs. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: Ciba-Geigy. Copyright NOVARTIS	Blade, ligule and sheath - line drawing	Ligule very short, 0.2-0.3 mm, with a conspicuous fringe of white hairs. Reproduced with permission from Hafliger E, Scholz H, 1981. Grass Weeds 2. Basle: Ciba-Geigy.	NOVARTIS
Title Whole plant - line drawing Caption a, Ligule, ventral view; b, part of spike; c, spikelet; d, flower; e, caryopsis. Copyright SEAMEO-BIOTROP	Whole plant - line drawing	a, Ligule, ventral view; b, part of spike; c, spikelet; d, flower; e, caryopsis.	SEAMEO-BIOTROP
Title Shoot Caption Shoot of C. dactylon showing three leaves per node. Copyright ©Chris Parker/Bristol, UK	Shoot	Shoot of C. dactylon showing three leaves per node.	©Chris Parker/Bristol, UK
Title Rhizome system Caption C. dactylon showing rhizome system (Botswana). Copyright ©Chris Parker/Bristol, UK	Rhizome system	C. dactylon showing rhizome system (Botswana).	©Chris Parker/Bristol, UK
Title Competition Caption C. dactylon competing with cotton in Egypt. Copyright ©Chris Parker/Bristol, UK	Competition	C. dactylon competing with cotton in Egypt.	©Chris Parker/Bristol, UK

Identity

Preferred Scientific Name

• Cynodon dactylon (L.) Pers.

Preferred Common Name

• Bermuda grass

Other Scientific Names

• Agrostis linearis Retz
• Agrostis stellata Willd.
• Capriola dactylon (L.) Kuntze
• Chloris maritime Trin.
• Chloris paytensis Steud.
• Cynodon affinis Caro & E.A.Sánchez
• Cynodon aristiglumis Caro & E.A.Sánchez
• Cynodon aristulatus Caro & E.A.Sánchez
• Cynodon decipiens Caro & E.A.Sánchez
• Cynodon distichloides Caro & E.A.Sánchez
• Cynodon erectus J.S. Presl. ex C.B. Presl.
• Cynodon glabratus Steud.
• Cynodon hirsutissimus (Litard. & Maire) Caro & E.A.Sánchez
• Cynodon iraquensis Caro
• Cynodon laeviglumis Caro & E.A.Sánchez
• Cynodon linearis Willd.
• Cynodon maritimus H.B.K.
• Cynodon mucronatus Caro & E.A.Sánchez
• Cynodon nitidus Caro & E.A.Sánchez
• Cynodon pascuus Nees
• Cynodon pedicellatus Caro
• Cynodon polevansii Stent
• Cynodon scabrifolius Caro
• Cynodon stellatus Willd.
• Cynodon tenuis Trin.
• Cynodon umbellatus (Lam.) Caro
• Cynosurus dactylon (L.) Pers.
• Cynosurus uniflorus Walter
• Digitaria dactylon (L.) Scop.
• Digitaria glumaepatula (Steud.) Miq.
• Digitaria glumipatula (Steud.) Miq.
• Digitaria linearis (L.) Pers.
• Digitaria maritima (Kunth) Spreng.
• Fibichia dactylon (L.) Beck
• Milium dactylon (L.) Moench.
• Panicum ambiguum (DC.) Le Turq.
• Panicum dactylon L.
• Panicum glumipatulum Steud.
• Panicum lineare L.
• Paspalum ambiguum DC.
• Paspalum dactylon (L.) Lam.
• Paspalum umbellatum Lam.
• Phleum dactylon (L.) Georgi
• Syntherisma linearis (L.) Nash
• Vilfa linearis (Retz.) P. Beauv.
• Vilfa stellata (Willd.) P. Beauv.

International Common Names

• English: Bahama grass; couch grass; devil grass; dog's tooth grass; quick grass; star grass
• Spanish: grama Bermuda; grama común; grama de Espana; gramilla; pasto bermuda; zacate de gallina
• French: chiendent; chiendent dactyle; gros chiendent; herbes-des-Bermudes; pied de poule
• Portuguese: capim-coastcross; capim-da-bermuda; capim-de-burro; grama; grama-bermuda; grama-seda; mate-me-embora

Local Common Names

• Angola: usila
• Argentina: chepica brava; grama bermuda; gramón; pasto de perro; pasto forestal; pata de perdiz; pie de gallina; tejedora; uña de gato
• Brazil: capim de burro; capim-bermuda; capim-fino; grama rasteteira; grama sao paulo; grama seda
• Cambodia: smao anchien
• Chile: pasto de galina
• Colombia: pasto Argentina; pasto ingles
• Cuba: grama; hierba de la Bermuda; hierba fina
• Dominican Republic: grama fina de Bermudas
• Egypt: negil
• El Salvador: barenillo; zacate de agujilla
• Fiji: balama grass; kabuta
• Germany: Echte-Hundsahn; Finger-Hundsahn
• Greece: agriada
• India: arugampul; doob; duba; hariali
• Indonesia: gigirintingan; jukut kakawatan; jukut raket
• Indonesia/Java: grintingan; hoe maneek; suket grinting
• Iran: chair
• Iraq: thayyel
• Israel: yableet matsui
• Italy: gramigna
• Japan: gyogishiba
• Lebanon: irk-en-najil; shirch-un unjil
• Malaysia: rumput minak
• Mexico: agrasia
• Morocco: mor-chiendent
• Myanmar: mye-sa-myet; mye-za-gyi
• Netherlands: hondsgrass
• New Zealand: Indian doab
• Pakistan: khabbal; talla
• Peru: grama dulce
• Philippines: babalut; galud-galud; kawad-kawaran; kulatai
• Puerto Rico: ala quete queda; pelo de brujas; pepe ortis; yerba Bermuda
• Saudi Arabia: nageel
• South Africa: gewone kweekgras
• Sri Lanka: aruham-pul; buha
• Sudan: nagila
• Suriname: tigriston
• Sweden: hundtandgraes
• Taiwan: gou-ya-gen
• Thailand: yah-phraek
• Turkey: kopek disi ayrigi
• USA/Hawaii: mahiki; manienie
• Venezuela: pasto pata de gallina
• Vietnam: cò chi'; co' ông
• Yugoslavia (Serbia and Montenegro): zubaca
• Zambia: kapinga

EPPO code

• CYNAR (Cynodon arcuatus)
• CYNDA (Cynodon dactylon)
• CYNMA (Cynodon maritimus)

Summary of Invasiveness

C. dactylon is a stoloniferous grass widely naturalized in tropical and subtropical regions of the world. This species is a C₄ grass included in the Global Compendium of Weeds (Randall, 2012) and it is listed as one of the most “serious” agricultural and environmental weeds in the world (Holm et al., 1977). It is a fast-growing grass that spreads by seeds and stolons and rapidly colonizes new areas and grows forming dense mats. As with many other African grasses, this species has the potential to alter ecosystem functions by altering fire regimes, hydrological cycles, biophysical dynamics, nutrients cycles, and community composition (D’Antonio and Vitousek, 1992). C. dactylon is very drought tolerant by virtue of rhizome survival through drought-induced dormancy over periods of up to 7 months. After dormancy, it has the ability to easily re-sprout from stolons and rooted runners. Plants also recover quickly after fire and can tolerate at least several weeks of deep flooding (Cook et al., 2005). Currently, C. dactylon is listed as invasive in many countries including Australia, Indonesia, Singapore, Cambodia, Vietnam, USA, Mexico, Costa Rica, Puerto Rico, Chile, Colombia, Uruguay, Argentina, Brazil and many islands in the Pacific Ocean such as Hawaii, Fiji, and French Polynesia among others (see distribution table for details). 

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Monocotyledonae
•                     Order: Cyperales
•                         Family: Poaceae
•                             Genus: Cynodon
•                                 Species: Cynodon dactylon

Notes on Taxonomy and Nomenclature

The name Cynodon dactylon is universally accepted for this common, widespread weed. It is highly variable and various subspecies have been distinguished (Harlan et al., 1970). It is normally tetraploid (2n=36) but diploid, triploid and pentaploid forms also occur (Rochecouste, 1962a; Thomas and Murray, 1978; Kissmann , 1991). A few other related species can also occur as weeds in some regions (see Similarities with Other Species).

Although it is not native to Bermuda, C. dactylon is an abundant invasive species there. It is presumed to have arrived in North America from Bermuda, resulting in its common name of Bermuda grass.
 

Description

C. dactylon is a perennial grass, with underground rhizomes and on the ground runners (Cabrera, 1968; Covas and Salvai, 1970). The runners spread horizontally and bear nodes with internodes of about 10 cm length. They may be flattened or cylindrical, mostly unhaired. Each node roots in the soil and produces short culms (tillers), up to 25 cm high, but develop into prostrate runners under less dense conditions. The almost unique character of the Cynodon genus of at least two and often three leaves at each node can be seen on the extended runners. This immediately distinguishes it from other perennial weeds with comparable growth pattern such as Panicum repens and Paspalum distichum (Perez and Labrada, 1985).

The rhizomes are mainly in the top 10 cm of the soil but may penetrate to a depth of 35 cm (Perez and Labrada, 1985; Phillips and Moaisi, 1993) .They may be twice as wide as the runners and this is one of the variable characters in populations (0.2-0.9 cm). Each node is covered by a white cataphyl. Runner or rhizome nodes may bear up to three viable buds.

Leaves have an alternate-distal pattern of distribution along the runners. Leaf blades are open up to the base, unhaired, similar or shorter than the length of the internode. The ligule is very short but with a conspicuous fringe of white hairs. Leaf blades are green to dull-green, from 1 to 15 cm depending on node, lanceolate, and forming almost a 90° angle with the leaf blade, finely parallel-ribbed on both surfaces, without a conspicuous midrib (Rosengurt et al., 1960). The width and pilosity of the weed blade may be used to distinguish populations of the weed (Oakley, 1999).

The inflorescence is supported on a culm up to 25 cm high and consists of a single whorl of 3-7 narrow racemes, each 3-8 cm long. Spikelets are 2-2.5 mm long in two rows, closely appressed to the rachis. Glumes are one-nerved, the lower almost as long as the spikelet, the upper half to three-quarters as long. The lemma is silky pubescent on the keel, palea glabrous. Caryopses are sub-eliptical, compressed and brownish, brilliant coloured (Kissmann, 1991).

The seedling has a hairy ligule, bearing 0.5 mm hairs. Pilosity increases as the seedling grows.

Plant Type

Distribution

C. dactylon is thought to have originated in Africa but now occurs worldwide in both tropical and subtropical regions including Asia, North, Central and South America, the Caribbean, and islands in the Pacific Ocean (see distribution table for details). It also spreads into temperate areas of Europe and North America but is limited by sensitivity to prolonged frost.

C. dactylon invades almost all kinds of crops and modified ecosystems, including urban areas and circulation paths (road and railroad tracks) in many regions. 

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The date of the initial introduction of C. dactylon into new habitats is unclear mainly because it has been repeatedly introduced in most countries where it is now present. In the USA, this species was introduced in 1807 and in Puerto Rico in 1876. Since 1943, many cultivars have been created and introduced in the USA, including among others the variety “Coastal” introduced in Georgia in 1943, “Alicia” introduced in Texas in 1967, “Grazer” introduced in Louisiana in 1985; and “Florakik” introduced in Florida in 1994 (Cook et al., 2005).

Risk of Introduction

C. dactylon has high potential for further spread in those areas where it is still absent. Ecophysiological and genetic traits coupled with both forms of propagation give this species a high score for success in almost any ecosystem.

Habitat

C. dactylon requires moderate warmth. It is tolerant of extremely high temperatures but is susceptible to hard or prolonged frost. It is especially predominant in subtropical conditions as a weed in both annual and perennial crops and in pastures, fallows and waste areas. It occurs under semi-arid and irrigated conditions on a wide range of soil types of varying pH and salinity.

Habitat List

Hosts/Species Affected

C. dactylon is treated by Holm et al. (1977) as the second most important weed in the world (after Cyperus rotundus), a status justified by its occurrence in virtually every tropical and subtropical country and in virtually every crop in those countries. In Holm et al. (1979) it is listed as a 'serious' or 'principal' weed in no less than 57 countries. A list of crops in which C. dactylon is, or could be, a problem weed would include virtually every crop of the tropics and subtropics and most temperate crops. The crops in which it is most commonly a major problem are those of the subtropics that are planted in wide rows, for example, cotton, sugarcane, tobacco, citrus, olive, deciduous fruit, forestry and ornamental species and many vegetables, but also some closer-planted but less competitive crops such as rice, lucerne, mixed lucerne and grass pastures, onion and jute (Labrada, 1994).

Host Plants and Other Plants Affected

Biology and Ecology

Genetics

The chromosome number reported for C. dactylon varies from 2n = 18 to 2n = 36 with diploid and polyploid populations (Cook et al., 2005).

Ramakrishan and Singh (1966) and Sarandon (1991) have found differences in total biomass and biomass partition according to the origin of the population. Sarandon (1991) points out that characters are highly heritable, which means that high genetic variability for biomass production and variable architecture allows an ample base for selection, which in most cases is induced by herbicides, mechanical control or forage production.

Reproductive Biology

C. dactylon is wind-pollinated and generally self-incompatible, suffering from inbreeding depression when genotypes are self-pollinated. Quantitative traits such as seed yield and forage yield can be dramatically negatively affected by inbreeding depression (Cook et al., 2005). In diploid populations, caryopses are formed after zygote formation. In polyploids, which are sterile, caryopses may be apomictic.

Physiology

This C4 plant (Kissmann, 1991) has high rates of accumulation under adequate irradiance, water and nutrient supply and may consume 75 kg of N, 20 kg of P and more than 1,500,000 litres of water for 5000 kg/ha of biomass dry matter (Fernandez , 1991). In the south of Santa Fe province, Argentina, a maximum biomass of 8000 kg/ha may be generated under a summer crop of maize or sunflower with >75 % located in the first 10 cm of the soil profile (Lombardo, 1973), whereas in Balcarce (Argentina) about 5000 kg/ha is commonly found in maize or sunflower stubble.

Phenology

A photoperiod of 13 hours induces flowering. Low night temperatures coupled with high diurnal temperatures induces blooming (Nir and Koller, 1976). A reduction in irradiance drastically decreases inflorescence production (Moreira, 1975). In North America, annual plants reproduce during spring and perennial plants reproduce all year long (USDA-NRCS, 2014).

Longevity

C. dactylon grows as both an annual and perennial grass. The annual growth-form becomes dormant and turns brown when nighttime temperatures fall below freezing or average daytime temperatures are below 10° C (Cook et al., 2005).

Activity Patterns

Seeds may be the route of invasion in weed-free fields through the faeces of cows (Rodriguez, personal communication). Rhizome biomass exhibits an annual cyclic pattern and, as with any perennial weed, low temperatures reduce biomass and viability is lost as a consequence of the consumption of materials due to respiration and maintenance. The digestibility of stocked material is severely decreased, implying a loss in forage quality (Vaz Martins, 1989). This is a character that has largely improved in cultivated varieties. Each node has a physiological self-governing structure in relation to the apex, but is highly dependent on substances from other plant parts. The mother plant determines the runner growth pattern on the soil surface according to the sugar-gibberellin balance (Montaldi 1970). Node disconnection may be caused by natural decay and cultivation and produces damage in the breakdown zone and changes in hormone and nutrient relationships. It is widely demonstrated that rhizome or runner fragmentation induces the activation of buds. The proportion of activated buds increases as the number of buds per segment decreases (Moreira, 1980; Kigel and Koller, 1985; Fernandez and Bedmar, 1992). The cultivation method is mainly responsible for vegetative propagation fragmentation. The higher the cultivation intensity, the smaller the segments produced (Kigel and Koller, 1985).

Population Size and Structure

This weed produces an enormous number of small seeds (0.25-0.30 mg), the viability and dormancy of which are highly variable according to genotype and the conditions when formed. The seed is important because it confers high genetic variability on the population.

Perez et al. (1995) recorded a very low germination rate. Uygur et al. (1985) obtained up to 15% germination at constant temperatures of 35-40°C, and 50% at temperatures alternating between 20 and 30°C. Moreira (1975) obtained up to 80% germination with the help of nitrate, chilling and alternating temperatures, and Elias (1986) recorded up to 96% germination from heavier samples of seed. Seeds remain viable in the soil for at least 2 years (Caixinhas et al., 1988). As a rule, cultivars have relatively high viability. Osmo-conditioning of Bermuda grass seeds with PEG followed by immediate sowing improved seed germination and seedling growth under saline conditions (Al-Humaid 2002).

The probability of emergence and successful establishment of C. dactylon decreases with the depth of the fragment, but increases with the weight of the node and internode (Perez et al., 1998). Growth from plants originated from a runner may exhibit a different biomass partition than that from plants originated from a rhizome (Fernandez, 1986). From sprouting onwards, weed growth is controlled mainly by temperature (optimum 25-30°C) and radiation, but also by humidity and soil fertility. The efficiency of carbohydrate reserve usage during sprout growth is highly dependent on temperature and the type of vegetative structure; it is maximum at 20°C and is higher for rhizomes than for stolons (Satorre et al., 1996). Runners and rhizome growth begins 30 days after growth but only if soil temperature is >15°C. Rates of 15 g/g/day have been recorded in Argentina (Lescano de Ríos, 1982).

Environmental Requirements

The annual cycle of the weed starts with bud reactivation by the end of winter. There is no innate dormancy and buds can activate if adequate temperature and humidity is available at any time of the year. Base temperature for bud sprouting ranging from 7.7°C, to 10°C (Bedmar 1991; Satorre 1996). New rhizomes are generated when temperatures exceed 15-20°C (Horowitz 1972). Aerial fractions have maximum dry matter content during the autumn and old rhizomes may have higher dry matter content than new rhizomes. C. dactylon tolerates a wide range of temperatures, especially very high temperatures in near-desert conditions. There is variation in tolerance to low temperatures and interactions between the time of exposure and relative humidity. Freezing point ranges from -2 to -3°C (Thomas 1969). Extreme temperatures were lethal to bud survival, but from 0 to 30°C, the rate of sprouting increased with increasing temperature.

Growth is favoured by medium-to-heavy, moist, well-drained soils but C. dactylon will also grow on acid and quite highly alkaline soils and the undisturbed rhizome system can survive flood conditions and drought (Holm et al., 1977). Runners allow C. dactylon to expand the exploration for new resources; avoid interference from other species; and propagate the most adapted genotype.

Air Temperature

Rainfall Regime

Soil Tolerances

Soil drainage

• free
• impeded

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• heavy
• light
• medium

Special soil tolerances

• saline
• shallow

Natural enemies

Notes on Natural Enemies

A wide range of natural enemies of C. dactylon have been recorded, including many that cause undesirable damage or disfigurement of C. dactylon lawns. They include a mycoplasma-like organism, which causes pale foliage and commonly occurs in weedy populations. Damage from natural enemies is rarely sufficient to provide useful control. Many of the species listed as natural enemies are better known as polyphagous pests of poaceous and other crops, while others such as Sipha maydis require evaluation before being considered as potential biological control agents attacking inflorescences and leaves in India (Labrada, 1994).

Means of Movement and Dispersal

C. dactylon is extremely efficient at propagation. Natural dispersal (non-biotic factors) such as water (flooding, irrigation) and agricutural practices (contaminated seed) and cattle husbandry propagate the seeds with released dormancy. Also cultivators, chisels and drilling equipment allow runners and rhizomes to be distributed away from a given source.

Pathway Vectors

Plant Trade

Impact Summary

Economic Impact

C. dactylon is treated by Holm et al. (1977) as the second most important weed in the world (after Cyperus rotundus), a status justified by its occurrence in virtually every tropical and sub-tropical country and in virtually every crop in those countries. In their later book (Holm et al., 1979), it is listed as a 'serious' or 'principal' weed in no less than 57 countries. It is especially vigorous and pre-dominant in crops with wide spacing (annual, such as cotton, and perennial, such as citrus) where it suffers a minimum of shading.

Holm et al. (1977) note that it is among the most serious weed species in sugarcane in 13 named countries, in cotton in eight, in maize in eight and in plantation crops and vineyards in 21. Other crops in which it is a serious problem include rice, groundnuts, tobacco, sugar-beet, banana, pawpaw, pineapple, sorghum and many vegetables. Its exact competitive potential has rarely been measured but it is generally regarded as a highly competitive weed. It has been listed among the top seven weed species in a world-wide review of weeds in sugarcane (Cepero and Rodriguez, 1983). There is some evidence for allelopathic effects on peach (Bengoa and Kogan, 1985) and on citrus (Horowitz, 1973) but it appears that such effects may depend on the crop species and local conditions.

In cotton, newly planted C. dactylon had little effect on cotton yield in the first season, but established stands with 75% or more ground cover reduced yields by 25-80% (Brown et al., 1985). In an earlier study, monetary loss from C. dactylon was estimated at $165/acre (ca $350/ha) (Kempen, 1984). The competitive effects of C. dactylon were generally less than those from the much taller grass weeds Sorghum halepense and Echinochloa crus-galli, but comparable with those from Solanum nigrum and species of Amaranthus. Another detailed study in the USA showed that cotton yields could be reduced by 40% by C. dactylon (Giraudo, 1992). A combination of C. dactylon and C. rotundus as dominant weeds left uncontrolled for 2 months reduced cotton yield by 90% in Brazil (Beltrao et al., 1978).

C. dactylon can act as host to a very wide range of organisms and is often implicated as an important alternate host of crop pests and diseases including, for example, bacterial leaf blight of rice (Li et al., 1985); covered smut of sorghum (Marley, 1995); stubborn disease of citrus (Carles, 1986); and others such as stripe disease of rice, barley yellow dwarf, lucerne dwarf, Rhizoctonia solani, Phyllachora sp. and Pratylenchus pratensis (see Holm et al., 1977 for further details). However, the extent to which C. dactylon has contributed to crop loss via these organisms has not been quantified.

C. dactylon may also cause cause poisoning in cows which pasture on the weed (Odriozola et al., 1998).

Losses to all weeds in a number of the crops in which C. dactylon is a major weed (for example, maize and cotton) are of the order of 10-15% and amount to many billions of US dollars. As a major component of the weed flora in these crops over a substantial proportion of their total area, it is likely that C. dactylon contributes at least 1 or 2% of this total monetary loss, i.e. many million dollars, to which could be added the time, effort and costs involved in manual weeding which is especially laborious in the case of this rhizomatous weed.

Environmental Impact

Hood and Naiman (2000) have compared the invasibility of riparian plant communities high on river banks with those on floodplain floors for four South African rivers. Analyses of abundant and significant riparian species showed that the floors have 3.1 times more exotic plants than the banks. The percentage of exotics ranges from 5 to 11% of total species richness for the banks, and from 20 to 30% for the floors. Species richness and percent exotics are negatively correlated for the banks, but not correlated for the floors. Authors claim that the most prominent exotic species are Lantana camara and C. dactylon, which are present on both banks and floors. Despite great differences in climate, species richness, and land use history, the percentages of exotic plants in three rivers in the Pacific Northwest of the USA and one river in southwestern France are similar to those in South Africa (24-30% versus 20-30%, respectively). Furthermore, the high proportions of exotic species in these riparian plant communities are comparable to those reported for vascular plant communities on islands. They so conclude that the macro-channel floor regions of the riparian zones of South African rivers are highly vulnerable to invasion by exotic vascular plants.

Threatened Species

Social Impact

The pollen of C. dactylon has been shown to produce allergy symptoms in asthmatics in Malaysia (Sam-ChoonKook et al., 1998) and Brazil (Kissmann, 1991). Purification and characterization of the isoallergen has been achieved by Chow (2003) .The role of C. dactylon in different religious ceremonies common to Bundelkhand region of Madhya Pradesh (India) has been studied by Dubey et al, (2000).

Auddy et al., (2003) have made extensive screenings of antioxidant activity of three Indian medicinal plants, including C. dactylon, traditionally used for the management of neurodegenerative diseases.

 

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Highly adaptable to different environments
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Highly mobile locally
• Has high reproductive potential
• Has propagules that can remain viable for more than one year

• Negatively impacts agriculture
• Negatively impacts human health
• Negatively impacts animal health
• Reduced native biodiversity

• Competition - monopolizing resources
• Competition - strangling
• Competition
• Pest and disease transmission

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect as a commodity contaminant
• Difficult/costly to control

Uses

C. dactylon is used in many countries  as a valuable pasture grass, a lawn grass, or an anti-erosion cover on bunds and embankments. In the USA, new cultivars with enhanced performance are periodically released (Eichhorn, 1996; Hanna et al., 1997). Mechanical and chemical methods of controlling weeds in C. dactylon tuffs have also been developed (Alvim et al., 1996; Johnson, 1997; Walker et al., 1998).

In areas where the soil is not suitable for growing crops such as maize or soyabean, C. dactylon has proved to be a forage resource for milk cattle. Its serious limitations for forage quality and production during winter may be much improved if suitable management (heavy instantaneous charge) allows the germination and establishment of naturalized species already existing in the soil bank such as Trifolium repens and Lolium multiflorum amongst others.

Somaclonal variation from C. dactylon germplasm can provide a source of resistance to Spodoptera frugiperda (Croughan et al., 1997).

Uses List

Animal feed, fodder, forage

• Fodder/animal feed
• Forage

Environmental

• Erosion control or dune stabilization

General

• Ornamental

Materials

• Poisonous to mammals

Medicinal, pharmaceutical

• Traditional/folklore

Similarities to Other Species/Conditions

Outside Africa, there is rarely confusion between C. dactylon and related species. However, in eastern Africa there is frequent confusion with C. nlemfuensis. This species is similar to C. dactylon in almost all respects but lacks below-ground rhizomes. Two larger, more vigorous species, which are also without rhizomes and can occur as weeds, are C. plectostachyus and C. aethiopicus. C. plectostachyus, which is often cultivated as giant star grass, is distinguished by the upper glume being very short, less than one-quarter the length of the spikelet; and C. aethiopicus differs mainly in being very robust and woody with multiple whorls of coloured racemes. For further details, descriptions and keys, see Clayton and Harlan (1970) and Clayton et al. (1974).

Prevention and Control

Cultural Control

Legumes or other cover crops are sometimes used for smothering C. dactylon since the weed does not tolerate deep shade. Vigorous crops and higher crop density may be important in reducing weed competition.

Mechanical Control

Traditional techniques of controlling C. dactylon rely very little on manual methods, as it easily survives shallow hoeing and positively thrives on mowing. However, the benefits of deep cultivation have been confirmed in Botswana and Zimbabwe where double ploughing, either after crop harvest or before the onset of the next season's rains, provided a high degree of control and was beneficial to crop yields (Phillips, 1993; Phillips and Moaisi, 1993; Mabasa et al., 1995). In these studies, conditions were dry and the rhizome systems not too deep, and standard ploughing depths of 10-15 cm were adequate to cause desiccation of the rhizomes. In contrast, Egamberdiev and Suleimanov (1985) found that double ploughing to 60 cm was needed, perhaps because wetter or cooler conditions meant that control had to be achieved by burial rather than desiccation. The main non-chemical approaches to control are deep tillage and shading/smothering crops.

Solarization

Solarization, using plastic sheets to raise soil temperatures, has proved highly effective in the control of C. dactylon in Egypt (Satour et al., 1991) and India (Nasr Esfahani, 1993; Anju-Kamra and Gaur, 1998). This technique has been also been recently tested in tomato orchards by Kumar et al (2003).

Chemical Control

EPTC may be used as a pre-plant incorporated treatment in soyabean and Phaseolus beans and, in combination with an herbicide such as dichlormid, in maize and sugarcane (Labrada, 1994). In each case a follow-up treatment with a post-emergent herbicide may be needed. In sugarcane, banana and plantain, glyphosate can be used successfully as a directed spray but care is needed to avoid contact with crop foliage. In sugarcane, dalapon may be a less hazardous choice for directed spraying, while an alternative for citrus and pineapple is the older soil-acting herbicide bromacil (Labrada, 1994).

Sulfometuron applied pre-emerge reduced C. dactylon ground cover from 73 to 93% in a succession-planted sugarcane crop in Louisiana, USA (Miller et al., 1998). Richard (1998) reported similar results with sulfometuron and also with thiazopyr. Richard et al. (2000) recently found that at-planting applications of clomazone mixed with either atrazine or sulfentrazone or imazapyr with atrazine were the only treatments evaluated that increased cane (7%) and sugar (9%) yields over the control of a weed community where C. dactylon was included.

Glufosinate-ammonium at maximum rate mixed with paraquat effectively controlled the weed under zero tillage in mustard and controlled C. dactylon under a non-crop situation (Das and Yaduraju, 2002.

Fluchloralin sprayed pre-sowing along with hand weeding gave effective control of C. dactylon in Cicer arietinum (Sesharee et al., 1996).

In other annual broad-leaved crops such as soyabeans, sunflower and potatoes, graminicides are selective and generally effective, with haloxyfop-methyl and quizalofop-ethyl having perhaps the highest activity, followed by fluazifop-butyl, and sethoxydim somewhat less effective (Kempen, 1983; Johnson and Talbert, 1985; Bedmar, 1997). Repeated applications may be needed for season-long control. The effects of different herbicidal combinations on weed management in soyabeans have been investigated by Jadhav et al. (2003).

The effects of addition of urea on herbicides have been tested by El-Quesni et al. (2000) in Egypt. Results suggest that glyphosate or fluazifop applied with urea gave an effective level of control of C. dactylon in July and August. Glyphosate formulations may have different behaviour, and have been studied by Martini et al. (2002): best results were obtained with 2.50 kg a.i./ha of potash glyphosate.

Imazapyr has proved selective in Pinus taeda and in Eucalyptus viminalis (Rivoir et al., 1987).

In tree and orchard crops, control of the actively growing weed before or after annual crops may be achieved using glyphosate (or the related sulphosate), which is effective at standard rates. The activity of glyphosate on C. dactylon is improved by a lower spray volume rate (Chivinge and Mare, 1991) and by various additives. However, a single application is unlikely to give permanent control and best results may be obtained by integration with tillage (Mabasa et al., 1995), smothering crops or ground covers: paraquat and glufosinate cause only temporary damage to the foliage.

The effects of most herbicides tend to vary according to the clone (biotype) or the population of C. dactylon being treated. Several studies have attempted to relate this to polyploidy level but little or no correlation has been detected for the activity of dalapon (Rochecouste, 1962b; Thomas and Murray, 1978) or sethoxydim (De Silva and Froud-Williams, 1992).

Biological Control

Drechslera cynodontis, Ustilago cynodontis, Puccinia cynodontis, and Fusarium poae as fungal pathogens and a specimen of Thripidae family were identified on C. dactylon and studied for potential use in biological control (Uygur, 2000).

Integrated Control

A single ploughing followed by glyphosate after a regrowth of C. dactylon may provide an effective and affordable control method to small-scale farmers (Abdullahi 2002).

Integrated weed management practices in spring planted sugarcane of coastal Orissa have been studied by Mishra et al. (2003). The effects of combined management techniques were studied in Italy by Caruso et al. (1997) who measured the effects of herbicides, and transplanting date of onions in the weed community structure. They found that crop growth rate increased during the first phases of the crop cycle, that the Shannon-Wiener diversity index showed a decreasing trend from the first to the third planting time for the herbicide treatment, and that the greatest mean influence on the diversity index was given by Amaranthus retroflexus, C. dactylon, Cyperus esculentus and Portulaca oleracea.

When designing a weed management program the following should be taken into account:

-The depletion of carbohydrates accumulated in the vegetative structures of the weed (sprouting promotion) coupled with the exposition of rhizomes and stolons to the soil surface in winter when low temperatures may kill buds to a greater extent.
-The design of a crop sequence, in which the initial crop must be highly competitive (such as sunflower or soyabean), to be sown early in the spring. The main target should be to avoid irradiance reaching this C₄ creeping weed.
-The use of herbicides such as glyphosate. These chemicals should be applied when the aerial/subterranean biomass ratio is at its highest value (at the end of favourable season), after the crop harvest in early autumn.

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Contributors

20/05/14 Updated by:

Julissa Rojas-Sandoval, Department of Botany-Smithsonian NMNH, Washington DC, USA

Pedro Acevedo-Rodríguez, Department of Botany-Smithsonian NMNH, Washington DC, USA

Distribution Maps