Spatial distribution of Lolium rigidum seedlings following seed dispersal by combine harvesters

This paper considers the relationships between the dispersal of seeds and the distribution pattern of an annual weed. A comparative study of seed dispersal by combine harvesters, with and without a straw chopper attached, was established using Lolium rigidum, a common weed in Mediterranean cereal crops. Seed dispersal distance was quantified and the relationships between dispersal and fine‐scale seedling distribution evaluated. Primary dispersal of L. rigidum seeds occurs in a very limited space around the parent plants, but the density of seed is low because most seeds do not fall from spikes spontaneously. In contrast, many seeds are spread by combine harvesters. In this study the maximum dispersal exceeded 18 m from established stands in cereal fields, although the modal distance was close to the origin. In addition, the action of the combine harvesters tended to accumulate L. rigidum seeds predominantly under the straw swath, with some lateral movement. This action could explain the fine‐scale banded pattern of L. rigidum in cereal fields. Although the treatment of straw by the standard and straw chopper combines differed, the resultant seed distribution showed few differences.     

Avena sterilis and Lolium rigidum infestations hamper the recovery of diverse arable weed communities

Intensification of agricultural practices has severely reduced weed diversity in arable fields, which affects the delivery of ecosystem services. However, in parallel, some species have benefited from intensive farming and have vastly increased their abundance, as is the case for Lolium rigidum and Avena sterilis in cereal fields. These highly competitive species severely reduce yields but can also compete with other weed species, and, when less intensive practices are applied, they might limit the recovery of weed diversity and the success of arable species reintroductions. A gradient of infestation was established in a winter wheat field in Catalonia (north‐eastern Spain) by sowing seeds of both species at three different densities to test their effects on the abundance, diversity and composition of the natural weed community. The emergence of seeds and the survival and biomass of transplanted seedlings of two rare species, Agrostemma githago and Vaccaria hispanica, were also evaluated. Avena sterilis and L. rigidum infestations reduced the diversity, abundance and biomass and changed the composition of the natural weed community, even at low infestation densities. Moreover, infestations of both species affected the overall performance of A. githago and V. hispanica. This study reveals that A. sterilis and L. rigidum are highly competitive and that their infestations might hamper the recovery of diverse weed communities. Their densities should be considered when selecting suitable sites for promoting diversity and reintroducing rare species.     

Multispectral classification of grass weeds and wheat (Triticum durum) using linear and nonparametric functional discriminant analysis and neural networks

Field studies were conducted to determine the potential of multispectral classification of late‐season grass weeds in wheat. Several classification techniques have been used to discriminate differences in reflectance between wheat and Avena sterilis, Phalaris brachystachys, Lolium rigidum and Polypogon monspeliensis in the 400–900 nm spectrum, and to evaluate the accuracy of performance for a spectral signature classification into the plant species or group to which it belongs. Fisher’s linear discriminant analysis, nonparametric functional discriminant analysis and several neural networks have been applied, either with a preliminary principal component analysis (PCA) or not and in different scenarios. Fisher’s linear discriminant analysis, feedforward neural networks and one‐layer neural network, all showed classification percentages between 90% and 100% with PCA. Generally, a preliminary computation of the most relevant principal components considerably improves the correct classification percentage. These results are promising because A. sterilis and L. rigidum, two of the most problematic, clearly patchy and expensive‐to‐control weeds in wheat, could be successfully discriminated from wheat in the 400–900 nm range. Our results suggest that mapping grass weed patches in wheat could be feasible with analysis of real‐time and high‐resolution satellite imagery acquired in mid‐May under these conditions.     

Spatial stability of Avena sterilis ssp. ludoviciana populations under annual applications of low rates of imazamethabenz

Long‐term experiments were conducted in two winter barley fields in central Spain to determine the spatial stability of Avena sterilis ssp. ludoviciana populations under annual applications of low rates of imazamethabenz herbicide. Weed density was sampled every year (over 5 years in the first field and over 3 years in the second) on the same grid locations prior to herbicide application. Although weed patches were stable in their location, weed density decreased in most of the years. In the first field, the populations decreased exponentially over the 5‐year period. The rates of population decline were dependent on the initial density of the population, being higher for the central core of the patches and lower for the low‐density areas. Under the conditions present in this experiment, it was possible to reduce heavy weed patches (up to 1200 seedlings m^?2) down to relatively safe levels (18 seedlings m^?2) in a period of 3 years using a density‐specific control programme, applying low rates of herbicides when weed densities were below a given level (1000 seedlings m^?2). However, under adverse environmental conditions, half rates of the herbicide failed to control the weed populations adequately. The stability of the location of patches of A. sterilis ssp. ludoviciana suggest that weed seedling distributions mapped in one year are good predictors of future seedling distributions. However, the actual densities established each year will depend on the control level achieved the previous year and the climatic conditions present during the establishment period.     

Assessing the opportunity for site-specific management of Avena sterilis in winter barley fields in Spain

The abundance of Avena sterilis in dryland barley fields was studied in four Spanish provinces. During two growing seasons, differential geopositioning system (DGPS)‐geo‐referenced A. sterilis infestations were obtained in 31 fields. The majority of the infestations were concentrated in few large but irregularly shaped patches, with a higher number of smaller and more regular patches accounting for a small proportion of the infestation. A multitude of very small and irregular patches completed the inventory. The implications of this spatial structure were studied. Site‐specific adjusted‐dose herbicide application offered 61–74% potential herbicide savings. However, given the low levels of infestation and the low economic returns obtained in most of the provinces, the most profitable strategy was generally no herbicide application. Site‐specific treatments were advantageous only in high‐returns systems. Because few large patches provided the majority of the infestation, zone‐specific treatments would be advisable, until such time that weed detection and site‐specific application technologies become more efficient.     

The effect of non-inversion tillage and light availability on dispersal and spatial growth of Cynodon dactylon

The invasive ability of Cynodon dactylon is dependent on self dispersal and on cultivation practices. Tillage can seriously change patch biomass and spatial structure, spreading vegetative propagules of the weed. The objectives of this study were: (i) to quantify the effect on non‐inversion tillage on dispersal, establishment and colonization of C. dactylon and (ii) to propose a simple model considering soil cultivation effects and light availability on spatial growth of weed patches. Two experiments were carried out, exploring different soils and environmental conditions. Spatial distribution of vegetative units differed when tillage was conducted with different non‐inversion implements and could be described by simple functions. A minimum patch biomass seems necessary before vegetative structures are vulnerable to movement by cultivation. Only a small proportion of the biomass dispersed from original patches was able to establish. However, simulation showed that the area colonized by C. dactylon mostly increased by means of tillage dispersal, both with and without crop competition, in one growing cycle. It appears sensible to consider changing cultivation practices to reduce weed dispersal and to use crop competition for light to create unsuitable habitats limiting weed colonization.     

Dispersal of Avena fatua and Avena sterilis patches by natural dissemination, soil tillage and combine harvesters

The dispersal of Avena spp. (A. fatua and A. sterilis) by natural dissemination and by agricultural operations was studied in four experiments conducted in Spain and Britain. Natural dispersal was very limited, with a maximum dispersal distance of 1.5 m. Dispersal was higher in the geographic direction that was downwind than in any of the other three geographic directions. Although plant movement was very small under no‐tillage, an annual patch displacement of 2–3 m in the tillage direction was observed under conventional soil tillage. Ploughing downhill resulted in much larger dispersal distances than ploughing uphill. In the crops studied, combine harvesters dispersed few Avena spp. seeds, because of the fact that the plants had shed most of their seeds (>90%) before harvest. The percentage of seeds available to be dispersed by the combine was dependent on the harvest time. Although combine harvesting may not contribute much to short‐distance dispersal, it may play an important role in long‐distance dispersal. In our studies, isolated plants were located up to 30 m from the original sources. This small proportion may have a significant effect on the distribution of the weed within a field, acting as foci for new patches.     

Comparison of growth,survivorship, seed production and shedding of eight weed species in a wheat crop in Western Australia

Development of integrated weed management strategies is dependent on a thorough knowledge of the demography of individual species. The current research established eight winter or summer weed species in a winter annual wheat cropping system at Wongan Hills, Western Australia, and investigated emergence of the first cohort of each species, survivorship, plant size, seed production and seed shedding over three years (2016–2019). The winter weeds Bromus diandrus and Lolium rigidum emerged at the same time as the wheat crop, and the initial cohort of marked plants had 100% survival to seed production in each year. By comparison, other winter weed species like Hordeum leporinum, Rumex hypogaeus, Sonchus oleraceus and Polygonum aviculare frequently emerged later than the crop and had a lower percentage of plants surviving to seed production. However, individual S. oleraceus and P. aviculare plants had the greatest seed production compared to other species. All winter weeds had variable patterns of seed shedding between years, with the exception of L. rigidum. Summer weed species emerged at the same time, but plants in the initial cohort of each species did not always survive to produce seed. The early emergence and high survivorship of B. diandrus indicates high competitive ability, but shedding commenced at a similar time to L. rigidum and harvest weed seed control may be a viable control method for this species.     

Comparison of sampling methodologies for site-specific management of Avena sterilis

The ability to manage weed infestations in a spatially precise manner requires efficient and accurate methods of mapping weed distributions. A study was conducted to compare four different ground-based methods for collecting georeferenced information on infestations of Avena sterilis in winter wheat and barley. Sampling was performed at harvest by scoring panicle density, either from the ground or from a combine, by counting the number of panicle contacts with a stick moved horizontally over the crop canopy by an observer walking through the field, and by sampling A. sterilis seed rain on the ground. No significant differences were observed among the populations estimated by the four methods. A partial budget analysis of the in-season costs and benefits of spraying patches using these methods showed that visual scoring from the combine was the most appropriate method for the creation of weed management maps to be used for patch spraying in the following season. As a large variety of spatial patterns may be found in fields, the recommended sampling method might be field-specific and optimality should be verified for general use.     

Multiple herbicide‐resistant Lolium rigidum (annual ryegrass) now dominates across the Western Australian grain belt

Lolium rigidum (annual or rigid ryegrass) is a widespread annual weed in cropping systems of southern Australia, and herbicide resistance in L. rigidum is a common problem in this region. In 2010, a random survey was conducted across the grain belt of Western Australia to determine the frequency of herbicide‐resistant L. rigidum populations and to compare this with the results of previous surveys in 1998 and 2003. During the survey, 466 cropping fields were visited, with a total of 362 L. rigidum populations collected. Screening of these populations with the herbicides commonly used for control of L. rigidum revealed that resistance to the ACCase‐ and ALS‐inhibiting herbicides was common, with 96% of populations having plants resistant to the ACCase herbicide diclofop‐methyl and 98% having plants resistant to the ALS herbicide sulfometuron. Resistance to another ACCase herbicide, clethodim, is increasing, with 65% of populations now containing resistant plants. Resistance to other herbicide modes of action was significantly lower, with 27% of populations containing plants with resistance to the pre‐emergent herbicide trifluralin, and glyphosate, atrazine and paraquat providing good control of most of the populations screened in this survey. Ninety five per cent of L. rigidum populations contained plants with resistance to at least two herbicide modes of action. These results demonstrate that resistance levels have increased dramatically for the ACCase‐ and ALS‐inhibiting herbicides since the last survey in 2003 (>95% vs. 70–90%); therefore, the use of a wide range of integrated weed management options are required to sustain these cropping systems in the future.     

Interactions between weeds of winter wheat under different fertilizer,cultivation and weed management treatments

Interactions between the weeds Bromussterilis L., Galium aparine L. and Papaver rhoeas L. were investigated over 3 years of continuous winter wheat (Triticum aestivum L.) across a factorial combination of organic and conventional fertilizer, and ploughing and hand-roguing contrasted with minimum tillage and herbicide. The species were sown separately and together at 50 seeds m^?2 per species at the start of the experiment in October 1989. In addition, there were weed-free and background-weed controls. Weed densities were monitored at roughly monthly intervals and crop yield recorded for three seasons. B. sterilis populations increased 10-fold under minimum tillage, but declined under ploughing. Densities of P. rhoeas remained largely low. G. aparine increased on the organically fertilized minimum-tillage plots, except where B. sterilis was present; the high densities of B. sterilis reduced the population size of G. aparine. Crop yield was influenced most strongly by the fertilizer treatment in the first season, but later the density of B. sterilis was by far the most important factor; the evidence of interactive effects of the different weed species on crop yield was weak at best. Interactive effects of arable weed species can be observed, but only at very high densities, and so are unlikely to be of widespread economic importance.     

Spatial correlation between weed species densities and soil properties

The spatial cross‐correlation between weed species densities and six soil properties within fields was analysed using cross‐semivariograms. The survey was carried out in three successive years in two fields. The most consistent relationship between weed species density (numbers m^?2) and soil properties was negative cross‐correlation between the density of Viola arvensis Murray and clay content. This correlation was found in both fields; however, the range of spatial dependence varied between fields. In one of the fields, the density of Lamium purpureum L. was positively cross‐correlated with the phosphorus content in the soil in all years. The density of Veronica spp. and Poa annua L. was negatively cross‐correlated with pH in all three years. Other spatial cross‐correlations that were found in this study were inconsistent over time or field site. The densities of some of the weed species were spatially cross‐correlated with more than one soil property. The results showed that the range of spatial dependence varied not only between fields, but also between weed species and soil properties, as well as between years. This study indicates that the weed pattern is field‐specific and that the spatial variation in soil properties within a field is one of several factors affecting weed patchiness.     

Molecular basis of multiple resistance to ACCase-inhibiting and ALS-inhibiting herbicides in Lolium rigidum

Herbicide resistance in Lolium rigidum is widespread across much of the agricultural land in Australia. As the incidence of herbicide resistance has increased, so has the incidence of multiple herbicide resistance. This reduces the herbicide options available for control of this weed. This study reports on the successful amplification and sequencing of the acetolactate synthase (ALS) gene of L. rigidum using primers designed from sequence information of related taxa. This enables, for the first time, the successful determination of a mutation in the ALS gene of this species that provides resistance to ALS‐inhibiting herbicides. This mutation causes amino acid substitution at Trp574 (numbering standardised to Arabidopsis thaliana) to Leu which had been reported to confer a high level of resistance against all classes of ALS inhibitor herbicides. In addition, multiple resistance to ALS‐inhibiting and acetyl‐coenzyme A carboxylase‐inhibiting herbicides is acquired through the independent accumulation of mutant alleles for the target sites. This may thus explain some of the irregular, mosaic resistance patterns that occur in this predominantly outcrossing species.     

A target-site mutation is present in a glyphosate-resistant Lolium rigidum population

Annual ryegrass (Lolium rigidum) is the only weed species to have evolved resistance to the broad‐spectrum herbicide glyphosate in Australia. A population that had failed to be controlled by glyphosate was collected from a vineyard in the Adelaide Hills region of South Australia. Dose–response experiments on this population (SLR 77) showed that it was glyphosate resistant, with an LD₅₀ that was 1.9–3.4 times higher than that of a susceptible population (VLR 1). The movement of radiolabelled glyphosate within SLR 77 plants showed that this population did not have the differential glyphosate translocation mechanism of resistance common to several other Australian glyphosate‐resistant populations. Subsequent analysis of shikimic acid accumulation within the plant after glyphosate treatment showed that this population accumulated significantly less shikimic acid than a susceptible population, but more than a glyphosate‐resistant population with the translocation mechanism, indicating the possible involvement of another mechanism of resistance. Sequencing of a portion of the SLR 77 5‐enolpyruvylshikimate‐3‐phosphate synthase gene was carried out and a mutation causing an amino acid change at position 106 from proline to threonine was identified. This mutation is likely to be responsible for glyphosate resistance in this population, as mutations in this position have been found to be responsible for glyphosate resistance in goosegrass (Eleusine indica) from Malaysia. This paper represents the first report of target‐site glyphosate resistance in L. rigidum and provides evidence that this species has at least two mechanisms of glyphosate resistance present in Australia.     

A survey of Lolium rigidum populations in citrus orchards: Factors explaining infestation levels

The presence of herbicide‐resistant Lolium rigidum in Mediterranean (Spanish) citrus orchards was reported in 2005 and it poses a serious threat to crop management. The main objective of this research was to investigate which components could be responsible for the persistence of annual ryegrass populations in Mediterranean mandarin and orange orchards. This is the first study regarding L. rigidum populations in Mediterranean citrus orchards. Surveys were conducted in 55 commercial citrus orchards in eastern Spain in 2013 by interviewing technicians who were working in cooperatives about crop management. The level of infestation by L. rigidum and the presence of harvester ants (Messor barbarus) then were estimated in the same orchards. The variables were subjected to a two‐dimensional analysis and both univariate and multivariate logistic regression models were fitted for each of the three L. rigidum density levels that had been established. The multivariate models showed the significant factors that were associated with various L. rigidum densities: (i) at a low density, the herbicides that were applied, the number of applications in 2013 and the type of irrigation (flood or drip); (ii) at a medium density, the presence of harvester ants; and (iii) at a high density, the herbicides that were applied in 2013. The results indicated that drip irrigation and one application of glyphosate mixed with other herbicides (or herbicides other than glyphosate) were associated with a lower L. rigidum density. The alternative management options that are presented here should help farmers to reduce weed problems in Mediterranean citrus orchards. Future research is required to better understand the presence of herbicide‐resistant populations, as well as the possible beneficial presence of granivorous ant species.     

Designing a sampling scheme to reveal correlations between weeds and soil properties at multiple spatial scales

Weeds tend to aggregate in patches within fields, and there is evidence that this is partly owing to variation in soil properties. Because the processes driving soil heterogeneity operate at various scales, the strength of the relations between soil properties and weed density would also be expected to be scale‐dependent. Quantifying these effects of scale on weed patch dynamics is essential to guide the design of discrete sampling protocols for mapping weed distribution. We developed a general method that uses novel within‐field nested sampling and residual maximum‐likelihood (reml ) estimation to explore scale‐dependent relations between weeds and soil properties. We validated the method using a case study of Alopecurus myosuroides in winter wheat. Using reml , we partitioned the variance and covariance into scale‐specific components and estimated the correlations between the weed counts and soil properties at each scale. We used variograms to quantify the spatial structure in the data and to map variables by kriging. Our methodology successfully captured the effect of scale on a number of edaphic drivers of weed patchiness. The overall Pearson correlations between A. myosuroides and soil organic matter and clay content were weak and masked the stronger correlations at >50 m. Knowing how the variance was partitioned across the spatial scales, we optimised the sampling design to focus sampling effort at those scales that contributed most to the total variance. The methods have the potential to guide patch spraying of weeds by identifying areas of the field that are vulnerable to weed establishment.     

Simulating the effects of weed spatial pattern and resolution of mapping and spraying on economics of site-specific management

The economic benefits of using site‐specific weed management (SSWM) are related to the proportion of the field that is weed‐infested, the number of weed patches and the spatial resolution of sampling and spraying technologies. In this paper we simulate different combinations of these factors using parameter values obtained for Avena sterilis ssp. ludoviciana growing in Spanish winter barley crops. The profitability of SSWM systems increased as the proportion of the field infested by this weed decreased and when patch distribution was more concentrated. Under most of the conditions tested, positive net returns for SSWM were obtained when the weed‐infested area was smaller than 30%. The highest net return occurred using a 12 m × 12 m mapping and spraying resolution. The critical parameter that determined the economic viability of patch mapping and spraying resolution was the technology costs. The site specific strategy was economically superior to the standard strategy (overall herbicide application) in most cases. However, the differential between the two strategies decreased when the number of patches and the resolution of mapping and spraying increased, such that the highest net returns were obtained with a single patch covering 14% of the field and using a 12‐m mapping and spraying resolution; whereas the worst net returns were obtained for all patch numbers when 64% of the field was infested and a 3‐m mapping and spraying resolution was used.     

Invasiveness of agronomic weed species in wheat in Western Australia

Weed seeds are introduced to agronomic systems naturally or through human-mediated seed dispersal, and introduced seeds have a high chance of being resistant to selective, in-crop herbicides. However, colonisation (invasion) rates for a weed species are usually much lower than rates of seed dispersal. The current research investigated colonisation of a winter annual wheat cropping system in Western Australia by a range of winter or summer annual weed species. The weed seeds were sown (at 100 seeds/m²) directly before seeding the crop in 2016 and allowed to grow in the following 3 years of wheat. Selective herbicides were not applied, to simulate growth of weed populations if the initial seed had been resistant to herbicide. Bromus diandrus, Hordeum leporinum, Rumex hypogaeus, Sonchus oleraceus, Polygonum aviculare, Lolium rigidum, Citrullus amarus and Tribulus terrestris colonised the crop, while Dactyloctenium radulans, Chloris truncata and Salsola australis failed to establish over 3 years. The most successful weed was B. diandrus, with a plant density of 1,170/m² by the third year and seed production of 67,740/m². The high density of B. diandrus reduced wheat density by 76% in the third year and reduced average yield by 36%. Lolium rigidum reduced average yield by 11%, and the other weed species did not affect crop yield. Further research is required on the invasiveness of these species in other regions, but it is clear that the spread of B. diandrus to new areas or the introduction of resistant B. diandrus seeds via contaminated grain should be avoided.     

Characterization of the germination and emergence response to temperature and soil moisture of Avena fatua and A. sterilis

Seeds of Avena fatua L. and )A. sterilis L. were germinated under a wide range of temperatures (5–30°C) and osmotic potentials (?25 to ?1400 KPa) in order to characterize their responses to these two environmental factors. Although both species behaved similarly at moderate temperatures, different responses were observed at the two extremes. )A. sterilis germinated and emerged in a higher proportion than A. fatua at temperatures below 10°C but the opposite was true at temperatures above 20°C. Although the rates of these two processes were similar in both species up to 18°C, above this temperature the germination and emergence of )A. sterilis was considerably delayed in comparison with that of A. fatua. The effect of decreasing osmotic potentials in reducing the germination was more pronounced in A. sterilis than in A. fatua. However, no differences were observed in the emergence responses of either species. The adaptative advantages of these characteristics and their relationship with the geographic distribution of the two species is discussed.     

Secondary dispersal, spatial dynamics and effects of herbicides on reproductive capacity of a recently introduced population of Bromus sterilis in an arable field

An arable field was subdivided and subjected to either deep inversion ploughing or non‐inversion cultivation after viable seeds of Bromus sterilis had been sown into oilseed rape stubble. After sowing in isolated plots distributed within the field, sequences of cropping treatments for the establishment of two successive winter wheat crops were applied. Each subfield was split into an uphill and a downhill direction for soil cultivation. The field had a 10° slope. In the season following seed introduction, 2.6% of the introduced seeds had successfully germinated and established in the non‐inversion cultivation regime, when no effective graminicide was applied. Ploughing eradicated B. sterilis. Using differential global positioning system (DGPS) mapping of the whole field population, emerged plants were observed up to 8.7 m (uphill treatment) and 21.3 m (downhill treatment) of their initial source. The median distance seeds were transported was 2.3 m uphill and 4.8 m downhill. Post‐emergence application of the herbicide propoxycarbazone slightly reduced weed density and seed weight, and almost halved weed seed production. Application of fenoxaprop‐P‐ethyl was followed by higher density of plants, tillers and seeds of B. sterilis. Seed viability was unaffected by herbicide use. Thus, in the second wheat crop following seed rain, the weed population was dispersed more widely in the field, such that 20–30% of seeds were dispersed more than 5 m distance from the first year's foci of infestation. The relevance of soil cultivation to secondary dispersal of B. sterilis is discussed.