Leptochloa fusca
(sprangletop)

Identity

Preferred Scientific Name

• Leptochloa fusca (L.) Kunth

Preferred Common Name

• sprangletop

Other Scientific Names

• Bromus polystachyus Kunth
• Centotheca malabarica (L.) Merr.
• Digitaria malabarica (L.) Roem. & Schult.
• Diplachne amboensis Roiv.
• Diplachne capensis (Nees) Nees
• Diplachne fusca (L.) P. Beauv. ex Roem. & Schult.
• Diplachne indica (Retz.) Spreng.
• Diplachne livida Nees
• Diplachne malabarica (L.) Merr.
• Diplachne pallida Hack.
• Diplachne parviflora (R.Br.) Benth.
• Diplachne reptatrix (L.) Druce
• Diplachne wahlbergii Roiv.
• Eragrostis procera (Roxb.) Steud.
• Festuca brownii F.Muell.
• Festuca fusca L.
• Festuca indica Retz.
• Festuca reptatrix L.
• Hemigymnia malabarica (L.) Henrard
• Leptochloa contracta (Retz.) Blatt. & McCann
• Leptochloa fascicularis (Lam.) A.Gray
• Leptochloa ginae Maire
• Leptochloa malabarica (L.) Veldkamp
• Leptochloa muelleri (Benth.) Stace
• Leptochloa neuroglossa Peter
• Leptochloa uninervia (J.Presl) Hitchc. & Chase
• Ottochloa malabarica (L.) Dandy
• Panicum malabaricum (L.) Merr.
• Poa contracta Retz.
• Poa fusca (L.) Desf.
• Poa malabarica L.
• Poa procera Roxb.
• Syntherisma malabarica (L.) Sw. ex Roem. & Schult.
• Tridens capensis Nees
• Triodia ambigua R.Br.
• Triodia capensis (Nees) T.Durand & Schinz
• Triodia formosana Honda
• Triodia livida (Nees) T.Durand & Schinz
• Triodia parviflora R.Br.
• Uralepis alba Steud.
• Uralepis capensis (Nees) Kunth
• Uralepis drummondii Steud.
• Uralepis fusca (L.) Steud.
• Uralepis livida (Nees) Steud.

International Common Names

• English: beetle grass; brown beetle grass; swamp grass
• Spanish: paja gris
• Chinese: shuang fu cao

Local Common Names

• USA: bearded sprangletop; littoral sprangletop; Malabar sprangletop; Mexican sprangletop

Summary of Invasiveness

L. fusca is a perennial weed with a global distribution. It is an aggressive species showing a competitive advantage in many situations due to its tolerance of saline and alkaline soils and its likely ability to fix nitrogen. It is commonly a serious weed of rice in many countries, and is of particular concern in Spanish rice fields. It is recorded as invasive on Hawaii and in the Chagos Archipeligo (as L. fusca ssp. uninervia) (PIER, 2014) and has been the subject of an ‘eradication action’ in Europe (Brunel et al., 2013).

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

Leptochloa fusca is the latest of a very large number of names applied to this species. Originally known as Festuca fusca by Linnaeus, it has been included in Poa, Digitaria and numerous other genera, including Diplachne.Peterson et al. (2012) make a strong case for use of the name Diplachne fusca and this is now used by USDA-ARS (2014), while most other authorities including the Plant List (2013) and USDA-NRCS (2014) continue to use Leptochloa fusca.

The Plant List (2013) recognizes three subspecies, fascicularis, muelleri and uninervia, in addition to ssp. fusca, with differing distributions as described by Peterson et al. (2012). These are all, at times, referred to as full species. The differing distributions are indicated to some degree in the distribution table, but for most purposes, the group is treated as a single species here.

The specific epithet ‘fusca’ means dark or dusky.

Description

Perennial, loosely tufted to rhizomatous. Culms erect or geniculate and rooting from lower nodes, up to 100 cm or more tall. Leaf sheaths glabrous; leaf blades tough, usually involute, 5-30(-50) × 0.15-0.3(-0.6) cm, adaxial surface scabrid, abaxial surface subglabrous; ligule 3-12 mm, acute. Inflorescence 15-25 cm, scabrid; racemes 3-28, indistinctly unilateral, 4-20 cm, straight, ascending or spreading, spikelets usually distant. Spikelets glaucous-green, subterete, 6-14 mm, florets 5-12; glumes keeled; lower glume lanceolate, 2-3 mm, acute; upper glume narrowly oblong, 3-4 mm, acute or mucronate; lemmas narrowly oblong, dorsally sub-rounded, lowest 4-5 mm, lower lateral veins pilose, entire or 2-dentate, midvein often produced into a short 0.3-1.6 mm awn; palea ciliolate along upper keels. Callus laterally pilose. Anthers 0.5-0.75(-2.5) mm. Caryopsis elliptic-oblong, 1.5-2.5 mm, dorso-ventrally flattened. Flowers from June to September (based on description of L. fusca ssp. fusca from Flora of China, 2014).

Light microscopic studies have established the existence of salt glands on leaf blades, with structure resembling that described for other halophytic genera within the Gramineae.

The subspecies fusca, ssp. muelleri, and ssp. uninervia, occurring in Australia, are separated as follows (AusGrass, 2014):

L. fusca ssp. muelleri: lowermost panicle branches not exserted at maturity; lemma often smoky white at maturity with a darker area surrounding the caryopsis.

L. fusca ssp. uninervia: lemma apex obtuse to truncate; lemma dark green or lead coloured; anthers almost always less than 0.7 mm long.

L. fusca ssp. fusca: lemma apex obtuse to acute or acuminate; lemma of various colours; anthers usually 0.5–2.5 mm long.

In USA, L. fusca ssp. fascicularis is distinguished from ssp. uninervia by its awned lemmas, 4-5 mm long (v. mucronate only, 1.5-2.5 mm long).

Distribution

The distribution indicated here is mostly for L. fusca in the broad sense. Each of the main subspecies, however, has a more restricted distribution which is indicated where the information is readily available (for example, from USDA-ARS, 2014). Further detail of the distribution of the individual subspecies can be gleaned from GBIF (2014), but has not always been included here.

A brief summary of native ranges for subspecies is as follows: L. fusca ssp. fusca is a polymorphic palaeotropical taxon native to Africa, Asia and Australasia; L. fusca ssp. muelleri is apparently restricted to Australia, known from much of the interior portions of eastern Australia, particularly the Northern Territory; L. fusca ssp. uninervia is native to and widespread in the Americas, from southern USA southwards; and L. fusca ssp. fascicularis is native almost throughout the temperate and tropical regions of the New World (Peterson et al., 2012; USDA-ARS, 2014).

L. fusca has been introduced to Hawaii, the Chagos archipelago, Midway Atoll, Tasmania and parts of Europe. In Spain, L. fusca ssp. uninervia and L. fusca ssp. fascicularis were found to be widely distributed in Valencia, increasing from a frequency of 5.3% in 2008 to 20.1% in 2010.

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

L. fusca ssp. uninervia and ssp. fascicularis are apparently recent introductions to Spain, where they were first recorded in 1990 (GBIF, 2014). In USA, L. fusca ssp. fascicularis has spread northwards and was first recorded in Michigan in 1920 (University of Michigan Herbarium, 2014). In Australia, L. fusca ssp. uninervis was first recognized in 1999 (Snow and Simon, 1999). In Italy, L. fusca ssp. fascicularis was first reported in 2000 and was thought to have been introduced from Spain (Romani and Tabacchi, 2000).

Introductions

Risk of Introduction

As a major weed of rice with a sizeable seed, L. fusca may be an occasional contaminant of unmilled rice and hence spread with imports of rice, but no instances of this happening can be quoted.

Habitat

In China, L. fusca it is a plant of shallow water, marshes and sometimes brackish ground (Flora of China, 2014). It is a salt-tolerant species and is known to excrete salt through glands on the leaves. Flora Zambesiaca (2014) describes it as growing in and beside shallow standing water, and in boggy, marshy, alluvial and black peaty soils and in wet sand, 0–1,280 m above sea level. It also describes it as a major weed on irrigation channels. In Spain, L. fusca ssp. uninervia is normally found around the edge of fields, while L. fusca ssp. fascicularis is found in the interior of flooded rice fields (Osca, 2013).

L. fusca has a relatively low tolerance to saline conditions at the seedling stage but tolerance increases during subsequent growth (Mahmood et al.,1995), and non-seedlings are tolerant of salinity, sodicity and alkalinity. Ola Hae et al. (2012) observed significantly reduced growth at 100 mM NaCl, but many other functions were normal and the plant was not severely damaged by 300 mM NaCl. In Egypt, fresh and dry weights increased with concentration of seawater between 12.5 and 25.0% (Ashour et al.,1997). Tolerance to NaCl appears to be associated with leaf extrusion and root efflux of both Na⁺ and Cl^-.

Ashok et al. (1996) reported that L. fusca showed remarkable tolerance to prolonged water stagnation of 30 days, due to enhanced root aerenchyma development and root growth, which enabled physiological processes and nutrient uptake to continue.

Habitat List

Hosts/Species Affected

L. fusca (mainly ssp. fasciculari) is a major weed of rice in a number of countries including USA, Cuba and Spain. L. fusca ssp. fusca is also problematic in rice in India and other countries. It can also occur in lucerne/alfalfa, tomatoes and turf.

Growth Stages

Biology and Ecology

Germination

Information on germination is often contradictory, perhaps reflecting differences between subspecies.

Fresh seed of L. fusca is normally dormant. Germination of dormant seed can be encouraged by chipping (Prota4U, 2013).

Baskin et al. (1999) suggested that L. fusca, as a semi-aquatic species, requires flooding for both dormancy loss and germination. Osca et al. (2011) found that seeds subjected to saturated or flooded soils had faster germination and development, resulting in heavier and bigger plants; however, L. fusca ssp. fascicularis and L. fusca ssp. uninervia do not germinate and emerge if the water level is maintained continuously above 10 cm (Osca et al., 2011), or when exposed to flooding (Mahmoud, 1997).

Myers and Morgan (1989) found that dormancy of L. fusca ssp. fusca seed was not broken by stratification, but did gradually break down during air-dry storage, indicating an after-ripening period of at least 1 year. Conversely, other authors have found stratification effective, perhaps reflecting differences between the subspecies. Osca et al. (2011) found that L. fusca ssp. fascicularis has a faster development than L. fusca ssp. uninervia when seeds had been stratified in saturated or flooded soil.

Studies by Mahmoud (1997) found similar germination rates in light and dark conditions and an optimum germination rate at an alternating temperature of 25/35°C. Salinity at 0.2% increased germination slightly, and although higher salinity levels decreased germination, L. fusca still achieved 30% germination at 1% salinity. A 2 mm soil covering was optimum for germination. Mahmoud (1997) found that germination was higher when seed had been stored moist at low temperatures than when stored dry and at room or high temperature.

Hong et al. (1995) found greatest germination rates (88% after four weeks) following dry rather than moist cold conditions, but confirmed the tendency for dormancy to reduce at lower temperatures. The germination rate slightly declined after storage of more than three months. Wet conditions and low temperatures were effective for inducing germination of seeds which had been stored under dry, room temperatures for four months. Germination rate was again favoured by an alternating temperature, with optimum germination rates achieved at 25/35°C for 10-14 hours. Hong et al. (1995) also confirmed germination at NaCl concentrations as high as 1.0%.

Physiology and Phenology

L. fusca is a C₄ plant, as indicated by its leaf anatomy, starch grain distribution, CO₂-compensation point and ¹³C:¹²C ratios (Yusuf and Malik, 1984).

Intensive studies of L. fusca in Pakistan suggest that there is a symbiotic relationship with Azoarcus bacteria, which may help with nitrogen fixation (Reinhold-Hurek et al., 1993b; Hurek et al., 1998; James, 2000). Field and greenhouse studies have shown that L. fusca may fix up to 26% of its nitrogen content (Malik et al., 1997). The mechanism by which this transfer occurs has still to be determined, and it may simply be via the relatively inefficient process of the death and mineralization of the bacteria.

The ability of Azoarcus sp. to infect L. fusca and rice was assessed in gnotobiotic culture and the genes involved in the infection process were investigated. The activity of two cellulolytic enzymes, exoglucanase and an endoglucanase, was detected in Azoarcus. Microaerobic conditions were required for nitrogen fixation in Azoarcus. Membrane stacks (termed diazosomes), with an iron protein uniformly distributed in the cytoplasm, were observed in spatial proximity to nitrogenase, leading to highly efficient coupling of respiration to nitrogen fixation ( Reinhold-Hurek et al., 1993b).

L. fusca ssp. fascicularis matures by June in Spain, thus allowing seeds to drop long before rice harvest (Osca, 2013).

Longevity

L. fusca is a biennial to perennial plant with a life span of two or more years (Ecocrop, 2014).

Nutrition

In India, L. fusca ssp. fusca responded significantly to nitrogen (Rao and Nayak, 2001) and to phosphorus (Abdullah et al., 2000).

As noted above, the ability of L. fusca to grow well on infertile soils is almost certainly due to its association with nitrogen fixing bacteria.

Associations

In Soetdoring Nature Reserve, Free State Province, South Africa, a brackish playa classified by Janecke et al. (2003) as an ‘Open Diplachne Pan’ is characterized by a Diplachne (=Leptochloa) fusca-Eragrostis bicolor community with an L. fusca subcommunity, and an Eragrostis bicolor-L. fusca subcommunity associated with it. L. fusca is dominant in the pan basin (Janecke et al., 2003). Also in South Africa an L. fusca-Acacia xanthophloea association has been described (Götze et al., 2003), as well as an L. fusca-Stipagrostidetea uniplumis association (Bezeidenhout et al., 1994).

As noted above, an association with nitrogen-fixing bacteria contributes greatly to its ability to grow well on infertile soils.

Environmental Requirements

L. fusca is a plant of tropical, sub-tropical and warm temperate climates.

Climate

Latitude/Altitude Ranges

Means of Movement and Dispersal

Natural Dispersal

Although L. fusca is perennial, with some short rhizomes, spread is very predominantly by seed. L. fusca seed can spread by water but has no adaptation to spread by wind.

Accidental Introduction

Osca (2013) emphasized the importance of spread via irrigation water. Taberner et al. (2011) noted the risk of spread by contaminated machinery.

Impact Summary

Economic Impact

L. fusca  ssp. uninervia and L. fusca ssp. fascicularis are significant weeds of rice in Spain (Osca, 2013).

In rice on saline soils in Kerala, India, L. fusca ssp. fusca was the most dominant weed species, occurring in approximately 85% of the sites surveyed (Vidya et al., 2004). It was also major weeds of rice in Cuba (Colon and Antigua, 1989), in Australia (McIntyre et al., 1989), in South Korea (Kang and Shim, 2002) and in Senegal (Haefele et al., 2000). L. fusca ssp. fascicularis is on the increase in Spain, where it represents a serious threat to rice crops, as it is found within the rice fields themselves (Osca, 2013).

In USA, L. fusca ssp. fascicularis interference durations of 63, 70 and 130 days after rice emergence reduced rice cv. Lemont grain yields by 11, 13 and 37% respectively. Interference durations of 21-56 days after emergence did not reduce grain yields (Carey et al., 1994).

Yields of drill-sown lowland rice cv. Lebonnet were reduced 9, 18, 20 and 36% by L. fusca ssp. fascicularis densities of 11, 22, 54 and 108 plants per square metre, respectively (Smith, 1983).

L. fusca ssp. uninervia was the most common weed species in the onion and tomato-green pepper plantation systems at Quíbor valley, Lara State, Venezuela (Martínez et al., 2003).

Environmental Impact

No reports of L. fusca effecting natural vegetation could be found. L. fusca can reduce salinity and sodicity in brackish and contaminated soils. A reduction in soil salinity of up to 37% in the surface (0-30 cm in depth) and 29.6% in the subsurface soil (30-60 cm in depth) was obtained after two years from transplantation of the grass in Egypt (Ashour et al., 2003). Farmers in Lodhran district, Pakistan Punjab, who had planted L. fusca for more than four years reported improved soil quality (Siddiqui et al., 2005).

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Long lived
• Has high reproductive potential
• Has propagules that can remain viable for more than one year
• Reproduces asexually

• Modification of nutrient regime

• Competition - monopolizing resources
• Competition - shading

• Difficult to identify/detect in the field
• Difficult/costly to control

Uses

Economic Value

Although L. fusca can grow in sites where other grasses do not flourish, opinions vary as to the value of it as forage. In India, planting of L. fusca on an alkali black clay soil was found suitable for green fodder and dry matter yields (Verma and Raghuwanshi, 2004). It is regarded as palatable but further data are needed both on the relative palatability to different animal species, as well as its nutritional qualities (Prota4U, 2013). In Australia, the liveweight of goats reduced when fed on L. fusca alone (Nawaz et al., 1993).

Yields of 19-40 tonnes per hectare per year have been obtained using various cutting regimes. An average protein content of 8.6% dry matter has been reported; it is also reported that increase in soil salinity is correlated with a decline in protein content, and with an increase in ash content (Prota 4U, 2013).

In India a Prosopis-Leptochloa system combines production with biological reclamation, and is an appropriate form of reclamation agroforestry for alkali lands (Singh, 1995).

Tawfik et al. (2013) suggested the potential of L. fusca as a biofuel in Egypt.

Similarities to Other Species/Conditions

In Flora of China (2104), L. chinensis and L. panicea are distinguished from L. fusca by being annual, with smaller, compressed spikelets, up to 4 mm long and with awnless lemmas.

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Cultural Control and Sanitary Measures

Gealy et al. (2013) recorded that the allelopathic rice cultivars PI 312777 and Taichung Native 1 (TN 1) provide significant suppression of  L. fusca ssp. fascicularis.

Physical/Mechanical Control

In Colombia, land levelling is recommended for control of  L. fusca ssp. fascicularis (Antigua, 1993).       

Movement Control

Machinery should be washed before moving between sites to reduce the risk of spreading L. fusca.

Chemical Control

Osca (2013) noted that L. fusca is susceptible to molinate and cyhalofop-butyl, and partially susceptible to propanil, but not susceptible to the more recently developed penoxsulam which is preferred for control of Echinochloa crus-galli. A switch in use to the latter herbicide is thought to have contributed to the intensification of L. fusca ssp. uninervia and ssp. fascicularis) in Spain.

Other herbicides reported effective in rice, mainly against L. fusca ssp. fascicularis, include thiobencarb and fenoxaprop + bentazone (Smith, 1988), granular chlomazone (Schulteis and Heier, 2003), dithiopyr, metolachlor, metolachlor + atrazine, pendimethalin and oxadiazon (McCarty et al., 1995). In addition, sequential applications of quinclorac and fenoxaprop, or propanil and sethoxydim (Stauber et al., 1991). In Cuba, pre-emergence thiobencarb or oxadiazon, and post-emergence propanil + thiobencarb have been recommended (Antigua, 1993). Isoxaben and atrazine treatments provide poor or inconsistent control (McCarty et al., 1995).

Glufosinate is effective in glufosinate-resistant (‘Liberty’) rice (Wheeler et al., 1998).

Nicosulfuron and fenoxaprop both gave excellent control of L. fusca ssp. fascicularis in turf (Grichar, 2011). In tomato in Peru, metribuzin and and pendimethaline were effective (Cerna and Rojas, 1979); and in lucerne/alfalfa, prodiamine (Fenderson et al., 1987).

Gaps in Knowledge/Research Needs

More research is needed on the to clarify the possibility of different germination behaviours and ecological requirements of the various subspecies.

References

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Contributors

29/03/17 Original text by: 

Chris Parker, Consultant, UK

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