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Cynodon

Cynodon is a of nine grass in the family , tribe Cynodonteae, with a base number of x = 9, native to the tropical regions of the . These plants are typically stoloniferous and/or rhizomatous, forming dense mats or turfs that spread vegetatively, and are valued for their adaptability to warm climates. Morphologically, of Cynodon feature culms that range from 4 to 100 cm tall, often branched, with flat, conduplicate, convolute, or involute leaf blades and sheaths that may be glabrous or pubescent. Inflorescences are terminal and digitate or subdigitate panicles comprising 1 to 20 spikelike branches, each bearing laterally compressed spikelets with one to three florets, where the glumes are shorter than the lemmas. The genus name derives from the Greek words for "" (kuon) and "" (odous), alluding to the toothed appearance of the rhizomes in some . Widely distributed now through human introduction, Cynodon species occur across subtropical and tropical zones globally, including in the United States, Pacific Islands, and parts of and . The most notable species, (Bermuda grass), is extensively used for lawns, turf, pastures, and due to its , rapid growth, and ability to withstand heavy traffic, though it is also considered an invasive weed in many non-native regions. Other species, such as C. aethiopicus (star grass) and C. nlemfuensis, serve similar and ornamental roles in . The genus includes accepted species like C. dactylon, C. aethiopicus, C. nlemfuensis, C. transvaalensis, C. incompletus, C. plectostachyus, and hybrids such as C. ×magennisii.

Description

Morphology

Plants in the genus Cynodon are perennial herbaceous grasses that typically form dense mats or turf through extensive horizontal growth via stolons and rhizomes. The culms are erect or ascending, ranging from 4 to 100 cm in height across species, and are often branched near the base; in C. dactylon, the most widespread species, culms are slender, wiry, and compressed, measuring 5-50 cm tall. Leaves consist of open sheaths without auricles and ligules that are either a fringe of hairs or a short membranous (0.2-0.5 mm long). Blade varies by species but is generally linear, flat, folded, or , with widths of 1-7 mm; for example, in C. dactylon, blades are 1-12 cm long and 1-4 mm wide, glabrous or sparsely pilose, with a subacute . The sheaths in C. dactylon are bearded at the mouth and otherwise glabrous. Inflorescences are terminal, digitate or subdigitate panicles composed of 2-20 spikelike branches, each 1-10 cm long and bearing two rows of solitary, overlapping spikelets. Spikelets are laterally compressed, 2-3 mm long, and contain 1-3 florets, with typically only the lowest floret fertile; glumes are membranous and keeled, shorter than the 3-veined lemmas, which may be hairy or winged in some species. In C. dactylon, the features 2-6 straight or arcuate racemes in a single whorl, each 1.5-7 cm long, with spikelets 2-2.7 mm that overlap by half to two-thirds their length. Rhizomes and stolons are robust and extensive, often reaching several meters in length, with roots developing at the nodes to support mat formation; in C. dactylon, rhizomes are slender and scaly. These structures contribute to the genus's ability to spread vegetatively. The base chromosome number for the genus is x = 9, with polyploidy common, resulting in diploid (2n = 18), tetraploid (2n = 36), and higher ploidy levels up to hexaploid (2n = 54) observed across species and accessions.

Reproduction and growth

Cynodon primarily reproduce asexually through vegetative via stolons, which are above-ground stems, and rhizomes, which are below-ground stems, enabling rapid clonal spread and the formation of dense mats that contribute to their persistence in various environments. This mode of allows plants to produce large numbers of genetically identical offspring efficiently, often dominating over in established populations. Sexual reproduction occurs via wind-pollinated flowers that produce , although seed viability and rates can vary significantly depending on environmental conditions and seed treatments such as . In , plants can produce substantial numbers of , supporting occasional despite the prevalence of clonal growth. As warm-season perennials, Cynodon species exhibit active growth in temperatures ranging from 15°C to 35°C, with optimal rates between 24°C and 37°C, and enter during cooler months when temperatures drop below 15°C. Establishment can occur from seeds, sprigs (vegetative cuttings), or plugs, achieving full coverage in 4 to 8 weeks under optimal warm, moist conditions. The lifecycle includes typically within 7 to 14 days at temperatures of 25°C to 30°C, followed by vegetative expansion and summer flowering that leads to seed set. Genetically, some Cynodon populations exhibit , an asexual seed formation process that results in uniform offspring genetically identical to the parent, enhancing clonal fidelity. Additionally, hybridization within the genus is possible, as seen in crosses between and Cynodon transvaalensis, which have produced sterile triploid cultivars used in turf applications.

Taxonomy

Etymology

The genus name Cynodon derives from the words kynós (κύων), meaning "," and odṓn (ὀδούς), meaning "," alluding to the sharp, hard, tooth-like scales on the rhizomes and stolons of plants in this . This etymological reference highlights a morphological feature reminiscent of a dog's tooth, a common in early botanical for descriptive purposes. The genus Cynodon was established by Louis Claude Marie Richard and formally published by Christiaan Hendrik Persoon in 1805, with designated as the . Prior to this, the type species had been described by in 1753 as Panicum dactylon, where the specific epithet dactylon originates from dáktylos (δάκτυλος), meaning "finger," in reference to the digitate arrangement of the racemes in the . Persoon's transfer of the species to the new genus consolidated its taxonomic placement within the family. Historically, species now classified under Cynodon were placed in other genera before the establishment of Cynodon, including Capriola proposed by in 1763, which was later rejected in favor of Cynodon under the Code of Nomenclature for algae, fungi, and . This consolidation reflects the evolving understanding of grass taxonomy in the late 18th and early 19th centuries, prioritizing morphological and phylogenetic coherence.

Classification and species

The genus Cynodon is classified within the family , subfamily , tribe , and subtribe Eleusininae. It was circumscribed by Christiaan Hendrik Persoon in 1805 in his Synopsis Plantarum. recognizes fifteen accepted in the genus as of 2024: including Cynodon ambiguus (Ohwi) P.M.Peterson, C. anisocarpus (Albr.) T.D.Macfarl., C. barberi Rang. & Tadul., C. convergens F.Muell., C. coursii A.Camus, C. dactylon (L.) Pers. (the ), C. incompletus Nees, C. nlemfuensis Vanderyst, C. plectostachyus (K.Schum.) Pilg., C. radiatus Roth, C. tenellus R.Br., and C. transvaalensis Burtt Davy, among others. Species distinctions arise primarily from differences in growth habit, architecture, and chromosome numbers, with most exhibiting levels from diploid (2n=18) to hexaploid (2n=54). For instance, C. dactylon is typically a low-growing, stoloniferous that forms dense through both stolons and rhizomes, often at tetraploid or higher ; in contrast, C. plectostachyus is a taller (culms 30–90 cm) with thick stolons and robust comprising 7–20 racemes in 2–7 whorls. The taxonomic history of Cynodon reflects ongoing refinements in grass systematics through the 19th and 20th centuries, driven by morphological and later molecular , leading to the exclusion of several originally placed in the . A notable example is Cynodon lagopoides (L.) R.Br., which was reclassified into the genus Aeluropus as A. lagopoides (L.) Trin. ex Thwaites based on differences in structure and adaptations. Current delimitations follow POWO (2024), incorporating phylogenetic analyses to resolve boundaries. Infrageneric divisions in Cynodon are informal and primarily based on levels and morphological variation rather than formal subgenera. Diploid species tend to be more localized and less aggressive, while polyploids show greater adaptability; the C. dactylon complex exemplifies this, encompassing the and its (e.g., C. dactylon subsp. dactylon and subsp. coursii), which vary in development and width but share stoloniferous habits.

Distribution and habitat

Native distribution

The genus Cynodon is native to warm temperate and tropical regions of the Eastern Hemisphere, encompassing parts of Africa, southern Europe, the Middle East, and southern Asia. Species within the genus originated primarily in African savannas and grasslands, with natural distributions shaped by pre-human dispersal mechanisms such as wind and animal migration. Cynodon dactylon, the most widespread species, is native across , including regions from southward to , as well as parts of southern such as . Cynodon plectostachyus is restricted to , occurring in countries like , , , , and , typically at elevations between 800 and 2,000 meters. Cynodon transvaalensis is endemic to , particularly , including provinces such as , , , and Northern Provinces. Prior to human influence, the genus was largely confined to latitudes between approximately 40°N and 30°S, thriving in diverse habitats from coastal plains to montane grasslands at elevations up to 2,000 meters. The evolutionary history of Cynodon traces back to origins within the , with diversification likely occurring through to arid and semi-arid environments over millions of years.

Introduced ranges and invasiveness

Cynodon dactylon, the primary species in the genus, was introduced to the during the colonial period, with the first documented arrival in the United States occurring in 1751 in , likely via contaminated or trade goods from . Subsequent intentional introductions for agricultural purposes, such as and establishment, facilitated its spread across the , where it has become naturalized in tropical and subtropical regions from southern to northern . In , particularly and , it arrived through similar colonial trade routes in the and has since established in disturbed habitats. The species now occupies a broad introduced range spanning approximately 45°N to 45°S , extending into temperate zones where mild winters allow persistence. Vectors of introduction include both deliberate human actions, such as seeding for in the early 1800s across southern U.S. states, and accidental transport via ships' hulls, contaminated hay, and machinery. Once established, C. dactylon rapidly colonizes disturbed soils, including roadsides, overgrazed pastures, and construction sites, due to its prolific and production, enabling quick dominance in open, sunny areas. This aggressive spread has led to its classification as invasive in multiple regions, where it outcompetes native vegetation by forming dense mats that reduce . In the United States, C. dactylon is designated a in several states, including and , where it invades grasslands and riparian zones, displacing native perennials like bunchgrasses. Similarly, in , it is recognized as an invasive environmental weed in all states, particularly in the south and west, where it alters native ecosystems by smothering plants. Control efforts often involve integrated methods, including selective herbicides like , though complete eradication is challenging due to its vegetative propagation. Currently, the covers over 12 million hectares in the U.S. for production, with cultivars further expanding its range into transitional zones.

Ecology

Environmental adaptations

Cynodon species, particularly C. dactylon, exhibit remarkable through a combination of morphological and physiological traits. The genus employs a photosynthetic pathway, which enhances water-use efficiency by concentrating CO₂ around , allowing sustained under reduced during water stress. Additionally, deep root systems, extending up to 2 meters in penetrable soils under drought conditions, enable access to subsoil moisture, while rhizomes facilitate survival during prolonged dry periods of up to seven months by entering and preserving carbohydrates. This allows C. dactylon to maintain growth at soil moisture levels below 50%, outperforming many C3 grasses in arid environments. Salinity resistance further underscores the adaptive prowess of Cynodon, with C. dactylon tolerating electrical (EC) up to 10 /m through mechanisms such as restricted sodium uptake and enhanced ion compartmentalization in roots. Ecotypes from saline habitats demonstrate high root proliferation and length under salt stress, supporting continued accumulation even at EC levels of 7-15 /m, beyond which growth slows significantly. adaptability spans optimal growth at 25-35°C, with mean daily temperatures above 24°C promoting vigorous development, while the plant endures brief frosts down to -10°C via protection, though foliage is killed below -2°C and regrows in spring. intolerance is pronounced, requiring at least 6 hours of full sun daily to prevent and die-off under medium to dense canopy cover. Soil versatility contributes to Cynodon's widespread success, thriving in ranges of 5-8 across sandy loams to clay loams, provided is adequate to avoid waterlogging, which induces . It performs poorly in highly acidic soils ( <5) or those with high aluminum saturation but adapts to low-fertility conditions with modest nutrient demands, requiring only 50-100 kg per annually for maintenance, though higher inputs boost productivity. is bolstered by tolerance, as rhizomes enable rapid resprouting post-burn without stand loss, and allelopathic root exudates—containing phenolics and acids—inhibit nearby competitors, reducing resource in disturbed habitats.

Interactions

Cynodon species, particularly C. dactylon, are wind-pollinated grasses that exhibit self-incompatibility. Seed dispersal occurs through endozoochory by mammals, as seeds are consumed and subsequently excreted, with enhanced germination observed in dung from livestock such as cattle and bison. These grasses form symbiotic associations with arbuscular mycorrhizal fungi (AMF), achieving infection rates of 95-96% in colonized , which enhance uptake, particularly , in nutrient-poor soils. AMF also improves overall growth and tolerance to like lead in contaminated environments. Additionally, Cynodon host nitrogen-fixing bacteria such as Azospirillum brasilense and related strains, which increase plant dry weight and total content by facilitating biological in the . These microbial partnerships contribute to the grass's adaptability in low-fertility habitats. As a forage resource, is heavily grazed by , including , goats, and sheep across African savannas, where it persists under intense pressure and supports hydrological improvements like reduced . However, it serves as an alternate host for various pests, including root-knot nematodes (Meloidogyne spp., such as M. marylandi), which form on roots and impair uptake. Viral pathogens like Cynodon chlorotic streak virus cause stunting and chlorotic streaking, while insects such as armyworms ( spp.) lead to defoliation and patchy damage in stands. In native savannas, Cynodon plays a key role by stabilizing shallow soils through its extensive network, significantly increasing resistance to and scour. In introduced ranges, however, its invasive growth forms dense monocultures that outcompete native forbs and reduce overall by dominating resources. This competitive edge is partly due to allelopathic effects mediated by , such as caffeic, ferulic, and p-coumaric acids, released from roots and leachates, which inhibit and growth of neighboring plants.

Cultivation and uses

Forage and pasture

Cynodon dactylon, commonly known as bermudagrass, serves as a primary warm-season grass in tropical and subtropical regions, providing reliable for such as and sheep. Other , such as C. aethiopicus (star grass) and C. nlemfuensis, are also valued for in tropical regions due to their productivity, palatability, and adaptability to various soils. Its vigorous growth and make it suitable for extensive grazing systems in areas with hot summers and mild winters. The species has a long history of use as forage, dating back to ancient times in Africa and India, where it was valued for sustaining livestock in arid environments. It was introduced to the United States in 1751, likely via contaminated hay shipments from colonial trade routes, and quickly adopted for hay and pasture production in the southeastern states. Nutritionally, C. dactylon offers moderate to high-quality forage with crude protein content typically ranging from 8% to 16%, depending on fertilization, maturity, and cultivar; levels can reach 16-20% under optimal nitrogen management. Digestible energy values generally fall between 2.2 and 2.6 Mcal/kg dry matter, supporting adequate weight gains in ruminants when grazed rotationally. The grass exhibits high palatability, preferred by cattle for its fine texture and digestibility. However, its oxalate content can bind dietary calcium in ruminants, potentially reducing absorption and leading to secondary deficiencies if not balanced with supplemental minerals. Effective management enhances productivity and nutritional quality. Grazing should begin when plants reach 5-10 cm in height, with stubble left at 3-5 cm to promote regrowth and prevent overgrazing. Nitrogen fertilization at rates of 200-400 kg N/ha annually, split into multiple applications, boosts yields and protein content; for instance, 80 kg N/ha post-grazing maintains vigor. Hybrids such as Tifton 85 outperform common varieties, yielding 20-30 tons of dry matter per hectare per year under intensive management with irrigation and fertilization, enabling stocking rates exceeding 1 animal unit per hectare during peak seasons.

Turfgrass and ornamental

Cynodon species, particularly C. dactylon (commonly known as bermudagrass), are extensively utilized in turfgrass applications for their exceptional wear tolerance and rapid recovery from damage, making them ideal for high-traffic areas such as golf courses, athletic fields, and lawns in warm climates. C. transvaalensis hybrids are also used in turf due to their fine texture and cold tolerance. These grasses form dense, fine-textured mats that withstand heavy foot traffic and mechanical stress, with cultivars like Tifway 419 serving as a longstanding standard for sports turf due to their vigorous spreading via stolons and rhizomes. In , bermudagrass is prevalent on NFL fields, where varieties such as Tifway 419 provide durability throughout the season, supporting play on natural grass surfaces in stadiums like those of the and . Establishment of Cynodon turf typically involves sodding for immediate coverage or for cost-effective , followed by regular mowing at heights of 1-3 cm to maintain a , low-growing sward suitable for recreational and ornamental purposes. Its inherent drought resistance, stemming from deep root systems, allows for reduced requirements compared to cool-season grasses, promoting water-efficient in arid or transitional zones. Ornamental applications include the use of select varieties for edging and borders, where their prostrate growth habit enhances aesthetic appeal in gardens and landscapes without invasive tendencies when properly managed. Maintenance of Cynodon turf focuses on preventing fungal diseases, such as dollar spot caused by Clarireedia homoeocarpa, which manifests as small, straw-colored lesions on leaves and can be controlled with targeted fungicide applications like or during humid conditions. The U.S. turfgrass generates approximately $2.2 billion in annual output (as of 2022), with Cynodon species contributing significantly through sod production and field installations, underscoring its economic importance in ornamental and recreational sectors. Historically, bermudagrass gained early adoption in the United States during the mid-19th century for stabilizing railroad roadbeds, such as along the Central of Georgia Railroad in , leveraging its soil-holding capabilities to prevent along tracks.

Potential risks

Cynodon species, particularly C. dactylon, can produce prussic acid (hydrocyanic acid) under stress conditions such as , , or mechanical damage, posing a risk to . This compound inhibits , leading to symptoms including rapid and , muscle tremors, staggering, and death within hours if ingested in sufficient quantities. A notable incident occurred in 2012 in , where 15 cattle died after consuming stressed Tifton 85 bermudagrass (C. dactylon × C. transvaalensis), confirmed by veterinary analysis to be due to prussic acid poisoning. As an in many regions, Cynodon dactylon displaces native vegetation through aggressive growth and competition for resources, thereby reducing local . In and riparian habitats, it forms dense monocultures that alter conditions and outcompete plants, leading to decreased habitat diversity for wildlife. For instance, in riparian areas, bermudagrass invasion has been documented to suppress establishment. Human exposure to Cynodon pollen can trigger allergic reactions, particularly seasonal (hay fever), affecting the with symptoms such as sneezing, , and itchy eyes. Bermuda grass pollen is a major in subtropical and temperate regions, with rates high among individuals in affected areas. Additionally, direct contact with the leaves may cause mechanical skin irritation due to abrasive silica phytoliths, resulting in characterized by redness and itching. In non-native arid and semi-arid regions, Cynodon cultivation demands substantial irrigation, exacerbating depletion and straining limited . Turfgrass like bermudagrass account for a significant portion of urban landscape water use, often requiring up to 1-2 inches of water per week during peak growth, which contributes to overall drawdown in water-scarce areas such as the .

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