CSIRO PUBLISHING

Crop & Pasture Science, 2019, 70, 917–925 Review

https://doi.org/10.1071/CP19261

Biology and management of Echinochloa colona and

E. crus-galli in the northern grain regions of Australia

A,C
Asad Shabbir , Bhagirath S. Chauhan B, and Michael J. Walsh A
A
Plant Breeding Institute, School of Life and Environmental Sciences, The University of Sydney,
Narrabri 2390, Australia.
B
Queensland Alliance for Agriculture and Food Sciences, The University of Queensland, Gatton,
Qld 4343, Australia.
C
Corresponding author. Email: [email protected]

Abstract. Echinochloa colona and E. crus-galli are two important annual grass weeds distributed throughout the summer
cropping regions of Australia. Both species are highly problematic weeds, responsible for yield losses of up to 50% in
summer grain crops. The success of Echinochloa species as weeds is attributed to their rapid growth, prolific seed
production, seed dormancy and adaptability to a wide range of environments. Importantly, E. colona has evolved resistance
to glyphosate in Australia, with resistant populations now widespread across the summer cropping regions. Fallow
management of E. colona with glyphosate alone is risky in terms of increasing the chance of resistance and highly
unsustainable; other control strategies (residual herbicides, strategic tillage, etc.) should be considered to complement
herbicides. This review provides a summary of current information on the biology, ecology and management of
Echinochloa species. The knowledge gaps and research opportunities identified will have pragmatic implications for
the management of these species in Australian grain cropping systems.

Additional keywords: barnyard grass, conservation cropping, weed biology, weed management.

Received 26 June 2019, accepted 2 October 2019, published online 20 November 2019

Introduction
there has been widespread evolution of herbicide resistance in
The northern cropping region of Australia, encompassing species such as Echinochloa colona (L.) Link (Charles et al.
Queensland and New South Wales (NSW), is one of the 2004; Cook et al. 2008).
important dryland production regions in Australia. Both winter Echinochloa colona and E. crus-galli (L.) P.Beauv. are
and summer crops are grown, predominantly wheat (Triticum problematic weed species in sorghum, mung bean (Vigna
aestivum L.) in winter and sorghum (Sorghum bicolor (L.) radiata (L.) R.Wilczek), maize (Zea mays L.), rice (Oryza
Moench) and cotton (Gossypium hirsutum L.) in summer sativa L.), cotton, and many other summer field crops of the
(ABARES 2018) with widespread use of summer fallows. grain-production regions of NSW and Queensland (Osten et al.
Yields of dryland winter crops rely on the stored soil water 2007; Pratley et al. 2008; Walker et al. 2010). These species
from the summer-dominant rainfall patterns. For effective soil negatively affect crop production through competition for soil
water storage, conservation-cropping systems based on reduced water and nutrients, either directly with summer crops or
tillage and residue retention are widely adopted across the indirectly by resource use during fallow phases preceding
northern Australian cropping region (Llewellyn et al. 2012). winter crops. Growers aim to control these species during
Conservation-cropping systems are more sustainable and summer crop and fallow phases. The annual cost for
reliable owing to improved soil health and soil-water controlling Echinochloa species and yield losses is estimated
retention. Over the last two decades, conservation-cropping at AU$14.6 million (Llewellyn et al. 2016).
systems have largely replaced conventional-tillage-based Echinochloa colona and E. crus-galli can be effectively
systems in Australia (D’Emden et al. 2006; Llewellyn et al. controlled with a range of pre- and post-emergent herbicides
2012). However, the reduced tillage practices and residue in summer crops and fallows (Fischer et al. 1966; Cook et al.
retention have resulted in a reliance on herbicides, especially 2008; Widderick et al. 2012). Glyphosate is a low-cost herbicide
glyphosate, for in-crop (glyphosate-tolerant cotton, G. hirsutum) that is heavily relied on for fallow and pre-seeding weed
and fallow weed control (D’Emden et al. 2006). This reliance on management. However, the evolution of glyphosate resistance
the same herbicide mode of action has led to the evolution of in E. colona has made control particularly difficult in glyphosate-
herbicide resistance in several weed populations that are now tolerant cotton and summer fallow systems (Walker et al. 2004;
widespread across the northern grain regions. Consequently, Thornby et al. 2013).

Journal compilation
CSIRO 2019 www.publish.csiro.au/journals/cp

918 Crop & Pasture Science A. Shabbir et al.

A better understanding of weed biology and ecology is critical E. crus-galli are awnless or have variable sizes of panicles
for development of effective weed-control techniques for and spikelets (Keenan et al. 2008; Pratley et al. 2008).
evolving cropping systems. Therefore, studying the biology of This phenotypic variation has implications for the correct
weeds in the context of a farming system can assist in identifying identification of Echinochloa species. In the rice-growing
new control techniques that can be complemented with regions of Australia, there is a zero-tolerance approach to
herbicides for effective weed management. In recent reviews, E. oryzoides (Ard.) Fritsch. in harvested rice grain (Pratley
some global perspectives on biology and management of et al. 2008); however, this species can be confused with large-
Echinochloa species have been provided (Bajwa et al. 2015; panicled forms of E. crus-galli.
Peerzada et al. 2016); however, more specific information on
these species with reference to Australian cropping systems is
lacking. The focus of this review is, therefore, to examine the History of introduction in Australia
available information on the biology, ecology and management The time of introduction of E. colona and E. crus-galli in
of E. colona and E. crus-galli with the aim of identifying research Australia is difficult to ascertain, mainly owing to their
priorities for developing alternative approaches to the control of obscure native range. Further, these species are cultivated as
these species in the northern cropping systems of Australia. fodder and forage species in different parts of the world
(ALA 2018; CABI 2018). In Australia, E. colona is generally
considered a naturalised species; however, because the origin is
Species description and distinction not well defined, some herbaria consider it to be native. The
Echinochloa colona, commonly known as awnless barnyard earliest herbarium record of E. crus-galli is of a collection in 1851
grass, jungle rice, swamp grass or Deccan grass, can have an in Queensland, probably as an escape from cultivation (ALA
erect or prostrate growth habit (Fig. 1). The purple bands on 2018). The first herbarium specimen of E. colona was collected in
leaves, reddish purple leaf sheaths, and awnless spikelets 1877, also from Queensland.
are important distinguishing characters of this species
(Table 1). Echinochloa crus-galli, commonly known as awned
barnyard grass, swamp barnyard grass, Japanese millet or Diversity and distribution in Australia
cockspur grass, is a robust tufted annual grass (Fig. 1), erect or In Australia, there are 19 species of Echinochloa, 10 native and
reclining at the base. The presence of awns, branched nine introduced. It is a complex genus with many intergrading
racemes, and longer leaf blades are some of the important forms, especially E. crus-galli, whose taxonomy is complicated,
characteristics that distinguish E. crus-galli from E. colona and this species has many indistinguishable taxa across Australia
(Table 1). Nonetheless, the size of panicle and presence or (Simon et al. 2009). E. crus-galli forms differ in habit, size of
absence of awns are not strict criteria for distinguishing spikelets and panicles, degree of awning, heading time, and
these species because several intermediate phenotypes of dormancy (Michael 1983; Pratley et al. 2008). Similarly,

Fig. 1. Young Echinochloa colona plant at its rosette stage showing typical purple bands on leaves (top
left); mature E. colona plant (bottom left); mature plant of E. crus-galli growing in an experimental field near
Narrabri, NSW (right), with a rosette growing in pot (inset right).
Biology and management of Echinochloa spp. Crop & Pasture Science 919

many forms of E. colona vary in height, growth habit, size of The two species are distributed throughout Australia;
panicles, and timing of heading (Keenan et al. 2008). E. colona occurs throughout the eastern and northern regions
Echinochloa colona and E. crus-galli are hexaploid species of Australia, whereas E. crus-galli is primarily found in the
with chromosome number 2n = 54 (Mulligan 1961; Yabuno southern and eastern regions (Fig. 2). The adoption of
1962; Michael 1983). Both species are self-compatible and conservation-cropping systems has favoured relatively small-
highly autogamous; however, some outcrossing (1.4%) seeded species such as E. colona, which has progressively
mediated by wind has been reported (Maun and Barrett 1986; climbed in rankings of weeds of importance in cotton
Nguyen 2015). production (Charles 1991). In surveys of cotton growers, this

Table 1. Biological attributes of Echinochloa colona and E. crus-galli as they occur in cropping systems

Biological attribute E. colona E. crus-galli Reference(s)

Leaf blade 3–25 cm long, 3–13 mm wide 5–50 cm long, 5–20 mm wide Holm et al. 1977
Panicles Appressed, 5–15 cm long Nodding, 5–20 cm long Holm et al. 1977
Spikelets 2–3 mm long, awnless 3–4 mm long, awned Holm et al. 1977
Germination cues Light and water Light and water Chauhan and Johnson 2009
Seed production 42 000 plant–1 40 000 plant–1 Holm et al. 1977
Seedbank size 24 750 seeds m–2 215 000 seeds m–2 Bagavathiannan et al. 2011; Widderick et al. 2012
Seedbank life 95% viability loss after 2 years 92% viability loss after 2 years Dawson and Bruns 1962; Walker et al. 2010
of burial at 0–2 cm depth of burial at 2.5 cm depth
Dormancy mechanism Innate and induced Innate Maun and Barrett 1986; Wu et al. 2004
Dormancy release 4 months 25% after 6 months Maun and Barrett 1986; Wu et al. 2004
Vegetative reproduction Yes No Holm et al. 1977
Time to reproductive stage 4–5 weeks after germination 4–5 weeks after germination Holm et al. 1977
Optimum growth temperature 25–288C 15–308C Maun and Barrett 1986; Chauhan and Johnson 2009

Distribution of Echinochloa spp. in Australia N

E. colona
E. crus-galli
Northern grain region

670 335 0 670 Kilometers

Fig. 2. Distribution patterns of Echinochloa colona and E. crus-galli across Australia (data Australasian Virtual Herbarium, https://avh.chah.
org.au).
920 Crop & Pasture Science A. Shabbir et al.

species was not listed as a troublesome weed of cotton in 1989 Dormancy patterns
(Charles 1991), but by 2014–15, it was ranked the second most Seeds of Echinochloa species exhibit high levels of dormancy
problematic weed after volunteer cotton (Manalil et al. 2017b). immediately after dispersal. In the northern cropping region,
This later survey found that E. colona was present in 67% of freshly produced seeds of E. colona have a very high degree
fields across the Australian cotton-production region (Manalil (100%) of primary dormancy, which is lost within 4 months of
et al. 2017b). soil burial (Walker et al. 2010). Various flushes of emergence are
By contrast, E. crus-galli occurrence has declined since reported in a single growing season, depending on soil water.
the initial 1989 survey of troublesome weeds of cotton, Like E. colona, fresh seeds of E. crus-galli are highly dormant,
present in 24% of surveyed fields (Charles 1991). The impact with only 1.4% germination at maturity (Rahn et al. 1968). Seeds
of E. crus-galli in cotton-production systems has declined of this species have an innate dormancy that is released after
substantially since this survey; it was not represented in the storage at room temperature for a period (12–14 months) that
top 15 weeds of 2010 (Werth et al. 2013) and was absent in varies between populations (Watanabe and Hirokawa 1974;
the most recent survey in 2014–15 (Manalil et al. 2017b). Miyahara 1975). A physical mechanism is likely involved,
because dormancy can be released by scarifying seeds with
Reproductive biology and life cycle acid, high temperature (40–508C) or puncturing with a scalpel
Echinochloa colona and E. crus-galli are fast growing, with both (Miyahara 1975; Sung et al. 1987). In Japan, E. crus-galli seeds,
species reportedly capable of flowering 4 weeks after after release from primary dormancy, may undergo secondary
germination, and with seed maturity and dispersal ~45 days dormancy in submerged soils or summer high temperatures
after germination (Peerzada et al. 2016). Echinochloa colona (>308C) (Miyahara 1975). This induced dormancy is released
typically establishes during the later stages of winter crops, to following low temperature from autumn to spring and during
begin reproductive development during late spring–early paddy flooding.
summer (Storrie et al. 2008). Seeds of Echinochloa species Understanding how dormancy affects the seasonal
are 2–3 mm in size, meaning that they are spread easily germination patterns of both species will allow better timing
through water, harvested grain, vehicles, agricultural of weed management. In Australia, detailed information on
machinery and birds (Maun and Barrett 1986). Individual dormancy-release mechanisms of different populations of
plants of E. colona and E. crus-galli can produce up to 42 000 Echinochloa species is lacking and needs further investigation
and 40 000 seeds, respectively (Holm et al. 1977; McGillion and to improve understanding of the ecology of these species.
Storrie 2006) (Table 1). Both species are adapted to flower under
a wide range of photoperiods ranging from 8- to 16-h daylength Emergence patterns and environmental cues
(Holm et al. 1977). Whereas E. crus-galli is a strict therophyte
In subtropical Australia, E. colona emerges in spring or early
and reproduces only through seeds, with no vegetative
summer when mean soil temperatures are >208C and soil-water
reproduction recorded (Maun and Barrett 1986), E. colona
levels are suitable (Walker et al. 2010). Under controlled
reproduces primarily through seeds but does have the ability
environments, seed germination can occur over a wide range
to multiply through lower nodes that contain adventitious roots
of alternating day/night temperatures from 208C/108C to
(Holm et al. 1977).
358C/258C (Chauhan and Johnson 2009; Mutti et al. 2019).
Echinochloa colona is a photoblastic species requiring light
Soil seedbanks and persistence to germinate. In the northern cropping region, it continues to
Knowledge of soil seedbanks and their persistence is important to germinate through summer and early autumn (Walker et al.
understanding the ecology and management of Echinochloa 2010), although the major flushes of emergence occur from
species across Australian cropping systems. Widderick et al. October to December. A minimum of 10 mm rainfall is
(2012) found an average 24 750 seeds m–2 of E. colona in the required for its emergence (Werth et al. 2017). Walker et al.
southern Darling Downs region of Queensland. This was larger (2010) studied the emergence dynamics of E. colona from
than the average E. crus-galli seedbank size of 6000 seeds m–2 different soil-burial depths in Australian dryland conditions
recorded in a survey of cropping fields in Arkansas, USA, but and found a general trend of decreasing emergence with
within the observed range of seedbank sizes (0–215 000 seeds increasing burial depth. Maximum emergence (28%) was
m 2) (Bagavathiannan et al. 2011). The soil seedbank life of recorded for shallow buried seeds (0–2 cm) and the lowest
E. colona can be relatively long (up to 6 years), depending on emergence (0.8%) for 10 cm burial depth over a 24-month
burial depth (Werth et al. 2013). Walker et al. (2010) studied the period. Emergence of E. colona is generally greatest from
persistence E. colona seeds when buried in soil at different moist and well-drained soils (Caton et al. 2004), but it can be
depths. After 24 months of burial, seed persistence declined severely restricted under flooded conditions (Estioko et al.
sharply from an initial 80% viability to 9% at 0–2 cm burial 2014).
depth and 10% at 5 cm, whereas 24% of seeds remained viable at No information is available on the seasonal emergence pattern
10 cm. This information suggests that in conservation-cropping of E. crus-galli in northern cropping systems of Australia. In
systems, where weed seeds remain at relatively shallow soil Ontario, Canada, maximum emergence of E. crus-galli occurred
depths (Swanton et al. 2000; Widderick et al. 2012), its seedbank at the start of summer (June) and continued intermittently until
can be depleted within 2–3 years. However, no information the end of the season (August) (Maun and Barrett 1986).
is available on the size of soil seedbanks and persistence of Optimum temperatures for E. crus-galli germination ranged
E. crus-galli in Australian cropping systems. from 158C to 308C (Maun and Barrett 1986). Similar to
Biology and management of Echinochloa spp. Crop & Pasture Science 921

E. colona, germination is highest for seeds near the soil surface. Current management options in conservation-cropping
Dawson and Bruns (1962) found that maximum emergence systems
(>70%) of E. crus-galli occurred from a depth of 1–2.5 cm Conservation-cropping systems are based on the practices of
and declined with increasing soil depth, with no emergence from reduced tillage and crop-residue retention aimed at improving
below 12 cm. soil quality and increasing water retention. The adoption of
conservation-cropping systems in Australia in the last
Weed–crop competition 30 years has been facilitated by herbicides, particularly
Some of the major competitive traits of Echinochloa species glyphosate, removing the need for cultivation and residue
include quick and prolific tiller production, profuse root growth burning for weed control; this has resulted in a dramatic
and a short life cycle. In Australia, data on crop-specific yield increase in reliance on herbicides for weed control (D’Emden
losses due to E. colona are not available but losses are likely to be and Llewellyn 2006). An unfortunate consequence of this
substantial. In a field competition study, Wu et al. (2010) used herbicide reliance has been the evolution of glyphosate
Japanese millet, E. esculenta (A.Braun) H.Scholz, a close mimic resistance in E. colona populations in the northern cropping
of E. crus-galli, and found that this model weed at a density of region (Cook et al. 2008; Thai et al. 2012).
80 plants m–2 can reduce sorghum yield by up to 1.6 t ha–1. In the
Philippines, a natural population (280 plants m–2) of E. colona Evolution of herbicide resistance
resulted in 76% yield losses in rice (Mercado and Talatala 1977). Growers across much of the northern grain region of Australia
In Arkansas, season-long competition with a high density have winter-cropping-based production systems that are reliant
(100–200 tillers m–2) of E. crus-galli reduced the grain yield on stored soil water from summer-dominant rainfall. A weed-
of rice by >40% (Smith 1974). free summer fallow is critical to the conservation of soil water for
Echinochloa crus-galli, when allowed to compete with cotton the subsequent winter crop, and to achieve this, glyphosate has been
for periods of 6, 9, 12 or 25 days after emergence, reduced the relied on for summer-fallow weed control. Frequent applications of
yield by 21%, 59%, 90% and 97%, respectively (Keeley and glyphosate during this phase have resulted in the evolution of
Thullen 1991). In a field study conducted in NSW, Kleinig and glyphosate resistance in E. colona populations as well as several
Noble (1968) found that in the absence of E. crus-galli other summer annual weed species (Walker et al. 2011; Werth et al.
competition and additional applied phosphorus (P), rice grain 2011). Glyphosate resistance in E. colona was first reported in
yield was significantly increased in response to nitrogen (N) Australia in 2007 in northern NSW (Cook et al. 2008), and
applied at 140 kg ha–1. However, with increasing density subsequently, several glyphosate-resistant populations have been
(172–236 to 366–548 tillers m–2) of E. crus-galli, the confirmed from the northern cropping region and other parts of
response to N was progressively reduced. Australia (Gaines et al. 2012; Thai et al. 2012).
The literature suggests that both Echinochloa species are There is a high potential for evolution of herbicide resistance
highly competitive and can significantly reduce the yield of many in populations of Echinochloa species as evidenced by the
crops around the world. However, there is a lack of information frequency of herbicide resistance in these species in global
on crop yield losses caused by these species in Australia. Detailed cropping systems (Heap 2018). Globally, populations of
studies are therefore needed to ascertain the crop-specific yield Echinochloa species have evolved resistance to 10 herbicide
losses caused by these species in northern cropping regions. modes of action (Table 2). Although herbicide resistance in

Table 2. Globally reported cases of herbicide resistance, classified by mode of action (MOA), in populations of Echinochloa colona and
E. crus-galli (from Heap 2018)

MOA Species Situation Countries

EPSP synthase inhibitors E. colona Fallow, cotton, rice, Argentina, Australia, United States
(glyphosate) watermelon
PSII inhibitors E. colona, Sorghum, corn Australia, Canada, Columbia, Costa Rica, El Salvador, France,
E. crus-galli Guatemala, Honduras, Iran, Panama, Poland, Spain, United States,
Venezuela
ACCase inhibitors E. colona, Rice, soybean Bolivia, Costa Rica, Egypt, Italy, Japan, China, Thailand, South
E. crus-galli Korea, United States, Venezuela
ALS inhibitors E. colona, Rice, corn Austria, Bolivia, Brazil, China, Egypt, France, Germany, Italy,
E. crus-galli Turkey, Ukraine, United States, Uruguay, Venezuela
Synthetic auxins E. colona, Rice Brazil, Columbia, United States
E. crus-galli
Very-long-chain fatty acid E. crus-galli Rice China, Philippines, Thailand
synthesis inhibitors
DOXP inhibitors E. crus-galli Rice United States
Cellulose inhibitors E. crus-galli Rice Brazil
Microtubule inhibitors E. crus-galli Orchards Bulgaria
Lipid inhibitors E. crus-galli Rice China, United States
922 Crop & Pasture Science A. Shabbir et al.

Australian populations of E. colona is currently restricted from 27% to 95%. Amaroo and Jarrah, two widely grown rice
primarily to glyphosate, the high potential for evolution of cultivars in Australia, were among the most suppressive
resistance has major implications for the management of this cultivars, with average suppression of weed root growth of
weed in summer fallows and crops in Australian summer- >90% compared with the other cultivars. Olofsdotter et al.
cropping systems (Werth et al. 2011; Manalil et al. 2017a). (1999) evaluated 111 rice cultivars for their ability to suppress
Several other herbicide options remain available (Iqbal et al. the growth of E. crus-galli in the field and later screened them in
2019), but the threat of resistance remains so there is a need the laboratory for their allelopathic potential. Cultivars differed
for additional and alternative weed-control techniques to significantly in their weed-tiller suppression (up to 77%) in the
complement herbicides. field and their root-length reduction (up to 63%) in the laboratory
with some correlation between both studies. Based on this, the
Crop competition authors indicated that allelopathy in combination with the
competitive ability of a rice cultivar determines the outcome
Several studies have demonstrated the potential for crop
of interference. No information is available on the allelopathic
competition to reduce the growth and development of
capability of crops against E. colona in Australian cropping
Echinochloa species. Chauhan (2013) found that a 75%
systems. However, it is expected that crop-specific allelopathic
shading treatment reduced the shoot biomass and seed
effects on E. colona would be similar to those of E. crus-galli.
production of E. colona 84% and 92%, respectively, and those
Weed-suppressive cultivars with improved allelopathic
of E. crus-galli by 70% and 76%.
interference may be created by conventional or modern
Reducing row spacing from 30 to 15 cm and increasing rice
breeding methods.
seeding rate from 30 to 50 kg ha–1 reduced densities of
Echinochloa species from 4.9 to 1.8 plant m–2 in direct-
seeded rice in India (Kaur and Singh 2014). Soybeans Strategic cultivation
(Glycine max (L.) Merr.) grown in 35-cm rows produced Cultivation is a mechanical weed-control technique that can be
46% higher yield and suppressed the growth of E. crus-galli used strategically (infrequently) to control relatively small-
by 80% compared with soybeans planted in wider 70-cm rows seeded weeds, such as Echinochloa species. In a study in the
(Kust and Smith 1969). southern Darling Downs region of Queensland, Widderick et al.
Some crop cultivars possess superior traits related to their (2014) evaluated the effect of different forms of tillage (harrow,
ability to access resources (such as light, water, nutrients), repeated chisel plough, and one-way disc plough) and stubble
making them more competitive against weeds (Manalil et al. cover (wheat, barley and sorghum) on the emergence and soil
2017a; Shabbir et al. 2017). For example, rice crop traits of high seedbank persistence of E. colona. The treatment with deep
biomass production, early canopy closure, longer stems, burial (>5 cm) and repeated tillage reduced emergence of
increased tillering capacity, and rapid leaf-area development E. colona by up to 57%. Emergence of E. colona was not
are all correlated with growth suppression of Echinochloa affected by the addition of different stubble treatments. In
species (Smith and Roy 1983; Ampong-Nyarko and De Detta another experiment, the same authors studied the effect of
1991; Bajwa et al. 2015). In a trial comparing 14 irrigated rice different tillage forms (no-till, harrow, Gyral (chisel), offset
cultivars for their competitive ability against E. colona, Fischer discs, and one-way disc) on vertical movement of small glass
et al. (1997) found that average yield losses in different cultivars beads mimicking E. colona seeds in the soil. Harrow and Gyral
ranged from 27% to 62%. In a field study, Mennan et al. (2012) treatments buried the smallest number (<2%) of beads, whereas
tested the competitiveness of five rice cultivars (Osmancık, offset disc and one-way disc treatments buried 27% and 66% of
Kızılırmak, Karadeniz, Koral and Negiş ) against E. crus-galli; beads below 5 cm in the soil. These studies indicate that E. colona
cv. Koral was ranked most suppressive with its ability to reduce emergence and seedbank may be reduced by tillage practices that
weed tiller production by 30% compared with the other cultivars. promote deep burial of seeds in the soil. By contrast, an earlier
In India, Kumar et al. (2013) found that Echinochloa species study had indicated that persistence increased with deep burial of
density and biomass were suppressed more when competing with seeds of E. colona (Walker et al. 2010). Therefore, the role of
cv. Prabhat than with cvv. Gautam and Krishna Hamsa. tillage and different types of tillage on burial and subsequent
There is an opportunity to integrate crop competition (narrow emergence of E. colona and E. crus-galli in northern cropping
row spacing, higher sowing rate, suppressive crop cultivars, regions requires further investigation.
planting times, etc.) with herbicides in control programs
that would likely improve the management of Echinochloa Research priorities for management of Echinochloa species
species. However, insufficient information is available on in conservation-cropping systems
crop competition effects on Echinochloa species in the
Detailed studies on effects of environmental factors on the eco-
northern cropping region and this warrants further research.
biological attributes (e.g. seed dormancy patterns, seedbank
dynamics, emergence patterns, persistence in soil) of different
Allelopathy populations of Echinochloa species have not been performed.
Allelopathy is a naturally occurring phenomenon of interference Such studies are needed to allow these species to be better
among species and can be exploited for weed management. Seal understood and managed across the conservation-cropping
et al. (2005) evaluated the allelopathic activity of 27 rice cultivars systems of Australia. For instance, detailed information on
on five rice weeds including E. crus-galli. Average root mechanisms of dormancy and its release in different
inhibition of all weeds by different Australian cultivars ranged populations of Echinochloa species is important for the
Biology and management of Echinochloa spp. Crop & Pasture Science 923

development of timing of management strategies against these Acknowledgements

species. We thank Grains Research Development Cooperation (GRDC US00084) for
Limited information is available on the effect of cultural supporting us to write this review paper. The authors declare that they have no
practices (e.g. planting time, row spacing, populations, conflict of interest.
cultivars) on the level of control of Echinochloa species in
Australian cropping systems. The integration of cultural
weed-control strategies with herbicides would likely provide
better control of these problematic species. Further, the References
development of non-chemical weed-control methods and ABARES (2018) Agricultural commodity statistics. Australian Bureau of
techniques will likely mitigate the problems associated with Agricultural and Resource Economics and Sciences, Canberra,
herbicide resistance in conservation-cropping systems. ACT. Available at: http://www.agriculture.gov.au/abares/research-
The literature contains contrasting information on the topics/aboutmyregion (accessed 14 April 2018).
effect of seed burial depth and tillage practices on the ALA (2018) Echinochloa P.Beauv. Atlas of Living Australia. Available at:
https://bie.ala.org.au/species/http%3A//id.biodiversity.org.au/node/apni/
persistence of E. colona. More information is required to
2898333#classification (accessed 13 March 2018).
improve understanding of the effect of strategic tillage on Ampong-Nyarko K, De Detta SK (1991) ‘A handbook for weed control in
persistence of Echinochloa species across conservation- rice.’ (International Rice Research Institute: Los Baños, The Philippines)
cropping systems. Bagavathiannan MV, Norsworthy JK, Smith KL, Burgos N (2011) Seedbank
The life histories of E. colona and E. crus-galli will need to size and emergence pattern of barnyardgrass (Echinochloa crus-galli) in
be studied in detail to understand their adaptation to different Arkansas. Weed Science 59, 359–365. doi:10.1614/WS-D-10-00149.1
cropping systems. For instance, the reason/s for the progressive Bajwa AA, Jabran K, Shahid M, Ali HH, Chauhan BS, Ehsanullah (2015)
decline in the presence of E. crus-galli in northern production Eco-biology and management of Echinochloa crus-galli. Crop
systems is not clear and requires further investigation. Protection 75, 151–162. doi:10.1016/j.cropro.2015.06.001
At present, there is not a clear understanding of the costs CABI (2018) Echinochloa crus-galli (barnyard grass). Centre for Agriculture
and Bioscience International, Invasive Species Compendium. Available
associated with the control of Echinochloa species in crops and
at: https://www.cabi.org/isc/datasheet/20367 (accessed 17 February
fallow. Research is needed to quantify the yield losses caused by 2018).
these species in major summer crops such as sorghum, Caton BP, Mortimer M, Hill JE (2004) ‘Weeds of rice in Asia.’ (International
mungbean, cotton and rice grown across conservation- Rice Research Institute: Los Baños, The Philippines)
cropping systems of Australia. Charles G (1991) A grower survey of weeds and herbicide use in the New
Site-specific weed-control techniques have gained interest for South Wales cotton industry. Australian Journal of Experimental
use in cropping systems over the last decade or so. Once weed Agriculture 31, 387–392. doi:10.1071/EA9910387
density is mapped, weed control can be done mechanically or Charles G, Taylor I, Roberts G (2004) The impact of the cotton farming
with herbicides. The role of emerging technologies such as system on weed succession: implications for herbicide resistance and
robotics and machine learning should be explored for adoption of an integrated weed management approach. In ‘Proceedings
14th Australian Weeds Conference’. Wagga Wagga, NSW. pp. 410–413.
managing Echinochloa species both in-crop and in fallows.
(Council of Australasian Weed Societies) Available at: http://www.
caws.org.au/awc/2004/awc200414101.pdf
Conclusions Chauhan BS (2013) Shade reduces growth and seed production of Echinochloa
colona, Echinochloa crus-galli, and Echinochloa glabrescens. Crop
Echinochloa species are highly problematic weeds infesting Protection 43, 241–245. doi:10.1016/j.cropro.2012.10.009
several crops and summer fallows and causing significant Chauhan BS, Johnson DE (2009) Seed germination ecology of Junglerice
losses to growers in the cropping systems of the northern (Echinochloa colona): a major weed of rice. Weed Science 57, 235–240.
region. Fast growth rate, wider photoperiod adaptability, doi:10.1614/WS-08-141.1
prolific seed production, and evolution of herbicide resistance Cook T, Storrie A, Moylan P, Adams B (2008) Field testing of glyphosate-
make these weeds very difficult to control in any cropping resistant awnless barnyard grass (Echinochloa colona) in northern
NSW. In ‘Proceedings 16th Australian Weeds Conference’. Cairns,
system. Fallow management of E. colona with glyphosate
Qld. (Eds RD van Klinken, VA Osten, FD Panetta, JC Scanlan)
alone is risky in terms of increasing the chance of resistance pp. 93. (Council of Australasian Weed Societies)
and highly unsustainable. Other control strategies should be D’Emden FH, Llewellyn RS (2006) No-tillage adoption decisions in
considered to complement herbicides. Applying crop southern Australian cropping and the role of weed management.
competition through manipulated agronomic practices Australian Journal of Experimental Agriculture 46, 563–569.
(e.g. altered row spacing and density, and competitive doi:10.1071/EA05025
cultivars) is an effective strategy for control of Echinochloa D’Emden FH, Llewellyn RS, Burton MP (2006) Adoption of conservation
species and should be considered in summer crops. Cultural tillage in Australian cropping regions: an application of duration
practices developed based on a better understanding of analysis. Technological Forecasting and Social Change 73, 630–647.
Echinochloa species biology and ecology, combined doi:10.1016/j.techfore.2005.07.003
Dawson JH, Bruns VF (1962) Emergence of barnyardgrass, green foxtail and
strategically with herbicides, would likely provide sustainable
yellow foxtail seedlings from various soil depths. Weeds 10, 136–139.
control of across diverse cropping systems in Australia. doi:10.2307/4040665
Estioko LP, Miro B, Johnson DE, Ismail AM, Baltazar AM, Merca FE (2014)
Differences in responses to flooding by germinating seeds of two
Conflicts of interest
contrasting rice cultivars and two species of economically important
The authors declare no conflicts of interest. grass weeds. AoB Plants 6, plu064. doi:10.1093/aobpla/plu064
924 Crop & Pasture Science A. Shabbir et al.

Fischer B, Swain DJ, Boerema EB (1966) Evaluation of three herbicides for Beauv. in water-seeded production systems. Crop Protection 41, 1–9.
the control of grasses in rices. Australian Journal of Experimental doi:10.1016/j.cropro.2012.04.027
Agriculture 6, 219–223. doi:10.1071/EA9660219 Mercado B, Talatala R (1977) Competitive ability of Echinochloa colonum
Fischer A, Ramírez HV, Lozano J (1997) Suppression of Junglerice L. against direct-seeded lowland rice. In ‘Proceedings 6th Asian-Pacific
[Echinochloa colona (L.) Link] by irrigated rice cultivars in Latin Weed Science Society Conference’. Indonesia. Vol. 1. pp. 161–165.
America. Agronomy Journal 89, 516–521. doi:10.2134/agronj1997. (Asian-Pacific Weed Science Society)
00021962008900030023x Michael PW (1983) Taxonomy and distribution of Echinochloa species with
Gaines TA, Cripps A, Powles SB (2012) Evolved resistance to glyphosate in special reference to their occurrence as weeds of rice. In ‘Proceedings
Junglerice (Echinochloa colona) from the tropical Ord River region Conference on Weed Control in Rice’. Los Baños, The Philippines.
in Australia. Weed Technology 26, 480–484. doi:10.1614/WT-D-12- pp. 291–306. (International Rice Research Institute: Los Baños, The
00029.1 Philippines)
Heap IM (2018) The International Survey of Herbicide Resistant Weeds. Miyahara M (1975) On dormancy of the seeds of Echinochloa crus-galli
Available at: http://www.weedscience.org/ (accessed 22 January 2018). Beauv. var. oryzicola Ohwi, a paddy field weed. Japan Agricultural
Holm LG, Plucknett DL, Pancho JV, Herberger JP (1977) ‘The world’s worst Research Quarterly 8, 194–198.
weeds: distribution and biology.’ (University Press Hawaii: Honolulu, Mulligan GA (1961) Chromosome numbers of Canadian weeds. III.
HI, USA) Canadian Journal of Botany 39, 1057–1066. doi:10.1139/b61-092
Iqbal N, Manalil S, Chauhan BS, Adkins SW (2019) Investigation of alternate Mutti NK, Mahajan G, Chauhan BS (2019) Seed germination ecology of
herbicides for effective weed management in glyphosate-tolerant cotton. glyphosate-resistant and glyphosate-susceptible biotypes of Echinochloa
Archives of Agronomy and Soil Science 65, 1885–1899. doi:10.1080/ colona L. (Link) in Australia. Crop & Pasture Science 70, 367–372.
03650340.2019.1579904 doi:10.1071/CP18444
Kaur S, Singh S (2014) Influence of crop density on weeds, growth and yield Nguyen TH (2015) Evolution and spread of glyphosate resistant barnyard
of direct-seeded rice. Indian Journal of Weed Science 46, 318–321. grass (Echinochloa colona (L.) Link) from Australia. PhD Thesis, The
Keeley PE, Thullen RJ (1991) Growth and interaction of barnyardgrass University of Adelaide, SA, Australia.
(Echinochloa crus-galli) with cotton (Gossypium hirsutum). Weed Olofsdotter M, Navarez D, Rebulanan M, Streibig J (1999) Weed suppressing
Science 39, 369–375. doi:10.1017/S0043174500073082 rice cultivars-does allelopathy play a role? Weed Research 39, 441–454.
Keenan MD, Walker S, Widderick M, Sindel B, Taylor I (2008) Barnyard doi:10.1046/j.1365-3180.1999.00159.x
grass responses to glyphosate—the importance of morphological traits. Osten VA, Walker SR, Storrie A, Widderick M, Moylan P, Robinson GR,
In ‘Proceedings 16th Australian Weeds Conference’. Cairns, Qld. p. 121. Galea K (2007) Survey of weed flora and management relative to
(Council of Australasian Weed Societies) Available at: http://www.caws. cropping practices in the north-eastern grain region of Australia.
org.au/awc/2008/awc200811211.pdf (accessed 22 May 2008) Australian Journal of Experimental Agriculture 47, 57–70. doi:10.1071/
Kleinig CR, Noble JC (1968) Competition between rice and barnyard grass EA05141
(Echinochloa). 1. The influence of weed density and nutrient supply in the Peerzada AM, Bajwa AA, Ali HH, Chauhan BS (2016) Biology, impact, and
field. Australian Journal of Experimental Agriculture and Animal management of Echinochloa colona (L.) Link. Crop Protection 83,
Husbandry 8, 358–363. doi:10.1071/EA9680358 56–66. doi:10.1016/j.cropro.2016.01.011
Kumar P, Singh Y, Singh UP (2013) Evaluation of cultivars and herbicides Pratley JE, Broster JC, Michael P (2008) Echinochloa spp. in Australian rice
for control of barnyard grass and nutsedge in boro rice. Indian Journal of fields species distribution and resistance status. Australian Journal of
Weed Science 45, 76–79. Agricultural Research 59, 639–645. doi:10.1071/AR07156
Kust CA, Smith RR (1969) Interaction of Linuron and row spacing for control Rahn EM, Sweet RD, Vengris J, Dunn S (1968) Life history studies as related
of yellow foxtail and barnyardgrass in soybeans. Weed Science 17, to weed control in the Northeast 5—barnyardgrass. Agricultural
489–491. doi:10.1017/S0043174500054655 Experiment Station University of Delaware, Newark, DE, USA.
Llewellyn RS, D’Emden FH, Kuehne G (2012) Extensive use of no-tillage in Seal AN, Pratley J, Haig T (2005) Evaluation of rice varieties for allelopathic
grain growing regions of Australia. Field Crops Research 132, 204–212. effects on Australian rice weeds—linking laboratory to field. In
doi:10.1016/j.fcr.2012.03.013 ‘Proceedings Fourth World Congress on Allelopathy’. 21–26 August
Llewellyn R, Ronning D, Ouzman J, Walker S, Mayfield A, Clarke M (2016) 2005, Charles Sturt University, Wagga Wagga, NSW. (Eds J Harper, M
Impact of weeds on Australian grain production: the cost of weeds to An, H Wu, J Kent) pp. 164–167. (International Allelopathy Society)
Australian grain growers and the adoption of weed management and Shabbir A, Ray P, Jabran K (2017) The future of weed science in the Asian
tillage practices. Report for GRDC, CSIRO Australia. Pacific Region—some perspectives for the next 50 Years. In
Manalil S, Coast O, Werth J, Chauhan BS (2017a) Weed management in ‘Commemorating 50 Years (1967–2017): 50th anniversary
cotton (Gossypium hirsutum L.) through weed-crop competition: A celebratory volume’. (Eds NR Chandrasena, AN Rao) pp. 111–124.
review. Crop Protection 95, 53–59. doi:10.1016/j.cropro.2016.08.008 (Asian-Pacific Weed Science Society, Indian Society of Weed Science,
Manalil S, Werth J, Jackson R, Chauhan BS, Preston C (2017b) An The Weed Science Society of Japan)
assessment of weed flora 14 years after the introduction of Simon BK, Weiller CM, Webster RD, Michael P (2009) Echinochloa (ms.)
glyphosate-tolerant cotton in Australia. Crop & Pasture Science 68, November 2009. In ‘Flora of Australia’. AusGrass2. Available at: http://
773–780. doi:10.1071/CP17116 ausgrass2.myspecies.info/node/3342 (accessed 16 August 2019)
Maun MA, Barrett SCH (1986) The biology of Canadian weeds: 77. Smith R (1974) Competition of barnyardgrass with rice cultivars. Weed
Echinochloa crus-galli (L.) Beauv. Canadian Journal of Plant Science 22, 423–426. doi:10.1017/S0043174500037930
Science 66, 739–759. doi:10.4141/cjps86-093 Smith J, Roy J (1983) Weeds of major economic importance in rice and yield
McGillion T, Storrie A (Eds) (2006) ‘Integrated weed management in losses due to weed competition. In ‘Proceedings Conference on Weed
Australian cropping systems: a training resource for farm advisors.’ Control in Rice’. Los Baños, The Philippines. pp. 19–36. (International
(Cooperative Research Centre for Australian Weed Management: Rice Research Institute: Los Baños, The Philipines)
Adelaide, S. Aust.) Available at: https://books.google.com.au/books? Storrie A, Cook T, Boutsalis P, Penberthy D, Moylan P (2008) Glyphosate
id=DkflAQAACAAJ resistance in awnless barnyard grass (Echinochloa colona (L.) Link) and
Mennan H, Ngouajio M, Sahin M, Isık D, Altop EK (2012) Competitiveness its implications for Australian farming systems. In ‘Proceedings 16th
of rice (Oryza sativa L.) cultivars against Echinochloa crus-galli (L.) Australian Weeds Conference’. Cairns, Qld. pp. 74–76. (Council of
Biology and management of Echinochloa spp. Crop & Pasture Science 925

Australasian Weed Societies) Available at: http://www.caws.org.au/ Werth J, Thornby D, Walker S (2011) Assessing weeds at risk of evolving
awc/2008/awc200810741.pdf (accessed 22 May 2008) glyphosate resistance in Australian sub-tropical glyphosate-resistant
Sung SS, Gerald RL, Hale MG (1987) Development and germination of cotton systems. Crop & Pasture Science 62, 1002–1009. doi:10.1071/
barnyardgrass (Echinochloa crus-galli) seeds. Weed Science 35, CP11201
211–215. doi:10.1017/S0043174500079078 Werth J, Boucher L, Thornby D, Walker S, Charles G (2013) Changes in
Swanton CJ, Shrestha A, Knezevic SZ, Roy RC, Ball-Coelho BR (2000) weed species since the introduction of glyphosate-resistant cotton.
Influence of tillage type on vertical weed seedbank distribution in a sandy Crop & Pasture Science 64, 791–798. doi:10.1071/CP13167
soil. Canadian Journal of Plant Science 80, 455–457. doi:10.4141/ Werth J, Keenan M, Thornby D, Bell K, Walker S (2017) Emergence of four
P99-020 weed species in response to rainfall and temperature. Weed Biology and
Thai HN, Malone J, Boutsalis P, Preston C (2012) Glyphosate resistance in Management 17, 29–35. doi:10.1111/wbm.12113
barnyard grass (Echinochloa colona). In ‘Proceedings 18th Australasian Widderick MJ, McLean AR, Walker SR (2012) Non-chemical options for
Weeds Conference’. Melbourne. pp. 237–240. (Council of Australasian managing the seed bank of glyphosate resistant weeds. In ‘Proceedings
Weed Societies) 18th Australasian Weeds Conference’. Melbourne. pp. 279–280.
Thornby D, Werth J, Walker S (2013) Managing glyphosate resistance in (Council of Australasian Weed Societies) Available at: http://
Australian cotton farming: modelling shows how to delay evolution and www.caws.org.au/awc/2012/awc201212791.pdf (accessed 11 October
maintain long-term population control. Crop & Pasture Science 64, 2012)
780–790. doi:10.1071/CP13109 Widderick M, Cook T, McLean A, Churchett J, Keenan M, Miller B,
Walker S, Widderick M, Storrie A, Osten V (2004) Preventing glyphosate Davidson B (2014) Improved management of key northern region
resistance in weeds of the northern grain region. In ‘Proceedings 14th weeds: diverse problems, diverse solutions. In ‘Proceedings 19th
Australian Weeds Conference’. Wagga Wagga, NSW. pp. 428–431. Australasian Weeds Conference’. Hobart, Tas. pp. 312–315. (Council
(Council of Australasian Weed Societies) Available at: http://www. of Australasian Weed Societies) Available at: http://www.caws.org.au/
caws.org.au/awc/2004/awc200414281.pdf (accessed 9 September 2004) awc/2014/awc201413121.pdf (accessed 4 September 2014)
Walker S, Wu H, Bell K (2010) Emergence and seed persistence of Wu H, Walker S, Osten V, Taylor I, Sindel B (2004) Emergence and
Echinochloa colona, Urochloa panicoides and Hibiscus trionum in persistence of barnyard grass (Echinochloa colona L. Link) and its
the sub-tropical environment of north-eastern Australia. Plant management options in sorghum. In ‘Proceedings 14th Australian
Protection Quarterly 25, 127–132. Weeds Conference’. Wagga Wagga, NSW. (Eds BM Sindel, SB
Walker S, Bell K, Robinson G, Widderick M (2011) Flaxleaf fleabane Johnson) pp. 538–541. (Council of Australasian Weed Societies)
(Conyza bonariensis) populations have developed glyphosate Wu H, Walker SR, Osten VA, Taylor IN, Robinson G (2010) Competition of
resistance in north-east Australian cropping fields. Crop Protection sorghum cultivars and densities with Japanese millet (Echinochloa
30, 311–317. doi:10.1016/j.cropro.2010.11.010 esculenta). Weed Biology and Management 10, 185–193. doi:10.1111/
Watanabe Y, Hirokawa F (1974) Ecological studies on the germination and j.1445-6664.2010.00383.x
emergence of annual weeds. 1. Effect of temperatures on the dormancy Yabuno T (1962) Cytotaxonomic studies on the two cultivated species and
breaking in seeds of Chenopodium album, Echinochloa crus-galli var. the wild relatives in the genus Echinochloa. Cytologia 27, 296–305.
praticola and Polygonum lapathifolium. Weed Research 17, 24–28. doi:10.1508/cytologia.27.296

www.publish.csiro.au/journals/cp