Testing solutions in grass-dominated landscapes: a review of current research

In this paper we review the experimental development of agri-environment measures for use on grasslands. Sward structure has been shown to have a strong influence on birds' ability to forage in grasslands, but the effects of food abundance on foraging behaviour are poorly understood and this hinders development of grassland conservation measures. The experiments described have a dual purpose: to investigate the foraging ecology of birds on grasslands and to test candidate management measures. Most of the work featured focuses on increasing invertebrate food resources during the summer by increasing habitat heterogeneity. We also identify important gaps in the habitats provided by existing or experimental measures, where similar dual-purpose experiments are required.

Leaving final-cut grass silage in situ overwinter as a seed resource for declining farmland birds

The loss of seed-rich wintering habitats has been a major contributory cause of farmland bird population declines in western Europe. Agricultural grasslands are particularly poor winter foraging habitats for granivorous birds, which have declined most in the pastoral farming regions of western Britain. We describe an experiment to test the utility of fertile ryegrass (Lolium) swards as a potentially rich source of winter seed for declining farmland birds. Four patches of final-cut grass silage were allowed to set seed and were left in situ overwinter. Half of each patch was lightly aftermath grazed in an attempt to increase the accessibility of the seed to foraging birds and reduce the perceived predation risk. Large numbers of yellowhammers (Emberiza citrinella) and reed buntings (E. schoeniclus) foraged on the seeded plots throughout the winter. They preferred to forage on ungrazed seeded plots, where the accumulation of senescent foliage resulted in a 14% average loss in silage yield in the following season. However, seed produced on the plots also led to sward regeneration, increasing subsequent yields on some plots. The technique offers clear benefits as a potential future agri-environment measure for declining granivorous birds, with wide applicability, but requires further development to minimise sward damage and costs to the farmer. Autumn grazing should reduce sward damage, but at the cost of reduced usage by buntings. Using the technique just prior to reseeding would be one way of avoiding any costs of sward damage.

Comparison of energy evaluation systems and a mechanistic model for milk production by dairy cattle offered fresh grass-based diets

Grass-based diets are of increasing social-economic importance in dairy cattle farming, but their low supply of glucogenic nutrients may limit the production of milk. Current evaluation systems that assess the energy supply and requirements are based on metabolisable energy (ME) or net energy (NE). These systems do not consider the characteristics of the energy delivering nutrients. In contrast, mechanistic models take into account the site of digestion, the type of nutrient absorbed and the type of nutrient required for production of milk constituents, and may therefore give a better prediction of supply and requirement of nutrients. The objective of the present study is to compare the ability of three energy evaluation systems, viz. the Dutch NE system, the agricultural and food research council (AFRC) ME system, and the feed into milk (FIM) ME system, and of a mechanistic model based on Dijkstra et al. [Simulation of digestion in cattle fed sugar cane: prediction of nutrient supply for milk production with locally available supplements. J. Agric. Sci., Cambridge 127, 247-60] and Mills et al. [A mechanistic model of whole-tract digestion and methanogenesis in the lactating dairy cow: model development, evaluation and application. J. Anim. Sci. 79, 1584-97] to predict the feed value of grass-based diets for milk production. The dataset for evaluation consists of 41 treatments of grass-based diets (at least 0.75 g ryegrass/g diet on DM basis). For each model, the predicted energy or nutrient supply, based on observed intake, was compared with predicted requirement based on observed performance. Assessment of the error of energy or nutrient supply relative to requirement is made by calculation of mean square prediction error (MSPE) and by concordance correlation coefficient (CCC). All energy evaluation systems predicted energy requirement to be lower (6-11%) than energy supply. The root MSPE (expressed as a proportion of the supply) was lowest for the mechanistic model (0.061), followed by the Dutch NE system (0.082), FIM ME system (0.097) and AFRCME system(0.118). For the energy evaluation systems, the error due to overall bias of prediction dominated the MSPE, whereas for the mechanistic model, proportionally 0.76 of MSPE was due to random variation. CCC analysis confirmed the higher accuracy and precision of the mechanistic model compared with energy evaluation systems. The error of prediction was positively related to grass protein content for the Dutch NE system, and was also positively related to grass DMI level for all models. In conclusion, current energy evaluation systems overestimate energy supply relative to energy requirement on grass-based diets for dairy cattle. The mechanistic model predicted glucogenic nutrients to limit performance of dairy cattle on grass-based diets, and proved to be more accurate and precise than the energy systems. The mechanistic model could be improved by allowing glucose maintenance and utilization requirements parameters to be variable. (C) 2007 Elsevier B.V. All rights reserved.

Effect of replacing grass silage with maize silage in the diet on bovine milk fatty acid composition

Even though extensive research has examined the role of nutrition on milk fat composition, there is less information on the impact of forages on milk fatty acid (FA) composition. In the current study, the effect of replacing grass silage (GS) with maize silage (MS) as part of a total mixed ration on animal performance and milk FA composition was examined using eight multiparous mid-lactation cows in a replicated 4 X 4 Latin square with 28-day experimental periods. Four treatments comprised the stepwise replacement of GS with MS (0, 160, 334 and 500 g/kg dry matter (DM)) in diets containing a 54:46 forage: concentrate ratio on a DM basis. Replacing GS with MS increased (P < 0.001) the DM intake, milk yield and milk protein content. Incremental replacement of GS with MS in the diet enhanced linearly (P < 0.001) the proportions of 6:0-14:0, decreased (P < 0.01) the 16:0 concentrations, but had no effect on the total milk fat saturated fatty acid content. Inclusion of MS altered the distribution of trans-18:1 isomers and enhanced (P < 0.05) total trans monounsaturated fatty acid and total conjugated linoleic acid content. Milk total n-3 polyunsaturated fatty acid (PUFA) content decreased with higher amounts of MS in the diet and n-6 PUFA concentration increased, leading to an elevated n-6: n-3 PUFA ratio. Despite some beneficial changes associated with the replacement of GS with MS, the overall effects on milk FA composition would not be expected to substantially improve long-term human health. However the role of forages on milk fat composition must also be balanced against the increases in total milk and protein yield on diets containing higher proportions of MS.

Enhancing pollinator biodiversity in intensive grasslands

Increased agricultural intensification has led to well-documented declines in the fauna and flora associated with intensive grasslands in the UK. We aimed to quantify the effectiveness of different field margin management strategies for putting bumblebee and butterfly biodiversity back into intensive grasslands. Using four intensive livestock farms in south-west England, we manipulated conventional management practices (addition of inorganic fertilizer, cutting frequency and height, and aftermath grazing) to generate seven grass-based treatments along a gradient of decreasing management intensity. We also tested two more interventionist treatments which introduced sown components into the sward: (i) a cereal, grass and legume mix, and (ii) a diverse conservation mix with kale, mixed cereals, linseed and legumes. These crop mixtures were intended to provide forage and structural resources for pollinators but were not intended to have agronomic value as livestock feed. Using a replicated block design, we monitored bumblebee and butterfly responses in 27 plots (10 x 50 m) in each farm from 2003 to 2006. Bumblebees were most abundant, species-rich and diverse in the sown treatments and virtually absent from the grass-based treatments. The diverse conservation mix treatment supported larger and more diverse bumblebee assemblages than the cereal, grass and legume mix treatment. The sown treatments, and the most extensively managed grass-based treatments, had the highest abundance, species richness and diversity of adult butterflies, whereas butterfly larvae were only found in the grass-based treatments. Bumblebee and butterfly assemblage structure was driven by floral abundance, floral richness, the availability of nectar resources, and sward structure. Only vegetation cover was correlated with butterfly larval abundance. Synthesis and applications. This study has identified management options in the margins of intensive grasslands which can enhance bumblebee and butterfly biodiversity. Extensification of conventional grass management by stopping fertilization, reducing cutting frequency and not grazing, benefits butterflies. However, to enhance bumblebees requires a more interventionist approach in the form of sowing flower-rich habitat. Both approaches are potentially suitable for adoption in agri-environment schemes in the UK and Europe.

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) canopy size on grass weed growth and grass weed seed return

Four experiments conducted over three seasons (2002-05) at the Crops Research Unit, University of Reading, investigated effects of canopy management of autumn sown oilseed rape (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) on competition with grass weeds. Emphasis was placed on the effect of the crop on the weeds. Rape canopy size was manipulated using sowing date, seed rate and the application of autumn fertilizer. Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds. were sown as indicative grass weeds. The effects of sowing date, seed rate and autumn nitrogen on crop competitive ability were correlated with rape biomass and fractional interception of photosynthetically active radiation (PAR) by the rape floral layer, to the extent that by spring there was good evidence of crop: weed replacement. An increase in seed rate up to the highest plant densities tested increased both rape biomass and competitiveness, e.g. in 2002/3, L. multiflorum head density was reduced from 539 to 245 heads/m(2) and spikelet density from 13 170 to 5960 spikelets/m(2) when rape plant density was increased from 16 to 81 plants/m(2). Spikelets/head of Lolium spp. was little affected by rape seed rate, but the length of heads of A. myosuroides was reduced by 9 % when plant density was increased from 29-51 plants/m(2). Autumn nitrogen increased rape biomass and reduced L. multiflorum head density (415 and 336 heads/m(2) without and with autumn nitrogen, respectively) and spikelet density (9990 and 8220 spikelets/m(2) without and with autumn nitrogen, respectively). The number of spikelets/head was not significantly affected by autumn nitrogen. Early sowing could increase biomass and competitiveness, but poor crop establishment sometimes overrode the effect. Where crop and weed establishment was similar for both sowing dates, a 2-week delay (i.e. early September to mid-September) increased L. multiflorum head density from 226 to 633 heads/m(2) and spikelet density from 5780 to 15 060 spikelets/m(2).

The influence of winter oilseed rape (Brassica napus ssp oleifera var. biennis) cultivar and grass genotype on the competitive balance between crop and grass weeds

Three experiments conducted over two years (2002-04) at the Crops Research Unit, University of Reading, investigated competition between autumn sown oilseed rape cultivars (Brassica napus L. ssp. oleifera var. biennis (DC.) Metzg.) and Lolium multiflorum Lam., L. x boucheanum Kunth and Alopecurus myosuroides Huds., sown as indicative grass weeds. Rape cultivar (cv.) had a substantial effect on grass weed seed return. Over the six cultivars tested, L. multiflorum spikelet production ranged from just under 400 spikelets/m(2) in the presence of cv. Winner to nearly 5800 in competition with cv. Lutin. Cultivar competitiveness was associated with high biomass, large dense floral layers and early stem extension. There was some evidence of differential competitive tolerance between rape cultivars. The results suggested that rape cultivars could be screened for competitiveness by measuring floral layer interception of photosynthetic active radiation. L. x boucheanum cultivars varied in ability to compete with rape. In the absence of inter-specific competition, spikelet density was similar for Aberecho and Polly (circa 31000 spikelets/m(2)) but when grown with rape Polly outyielded Aberecho (i.e. 12 090 and 7990 spikelets/m(2) respectively).

The value of sown grass margins for enhancing soil macrofaunal biodiversity in arable systems

The presence of a grass strip was found to be beneficial to soil macrofauna, increasing the species densities and abundances of earthworms, woodlice and staphylinid beetles. The biodiversity of the three main feeding groups - predators, soil ingesters and litter consumers - was also significantly higher in the grass strips than in the field edges without strips, indicating that establishment of grassy margins in arable fields may enhance ecosystem services such as soil fertility and pest control. The grass strip habitat contained a large number of species of soil macrofauna, being second only to hedgerow habitat, with 10% of the total species list for the farm found only within the margins. Of the rare species recorded on the farm, five of the nine were from the grass strips, four of which were found only there. This study shows that establishing grassy strips in the margins of arable fields increases the biodiversity of the soil macrofauna, both within fields (alpha diversity) and across the farm (beta diversity). (C) 2008 Elsevier B.V. All rights reserved.

Factors affecting biomass production and nutritive value of Calliandra calothyrsus leaf as fodder for ruminants

Calliandra calothyrsus is a tree legume native to Mexico and Central America. The species has attracted considerable attention for its capacity to produce both fuelwood and foliage for either green manure or fodder. Its high content of proanthocyanidins (condensed tannins) and associated low digestibility has, however, limited its use as a feed for ruminants, and there is also a widespread perception that wilting the leaves further reduces their nutritive value. Nevertheless, there has been increasing uptake of calliandra as fodder in certain regions, notably the Central Highlands of Kenya. The present study, conducted in Embu, Kenya, investigated effects of provenance, wilting, cutting frequency and seasonal variation both in the laboratory (in vitro digestibility, crude protein, neutral detergent fibre, extractable and bound proanthocyanidins) and in on-station animal production trials with growing lambs and lactating goats. The local Kenyan landrace of calliandra (Embu) and a closely-related Guatemalan provenance (Patulul) were found to be significantly different, and superior, to a provenance from Nicaragua (San Ramon) in most of the laboratory traits measured, as well as in animal production and feed efficiency. Cutting frequency had no important effect on quality; and although all quality traits displayed seasonal variation there was little discernible pattern to this variation. Wilting had a much less negative effect than expected, and for lambs fed calliandra as a supplement to a low quality basal feed (maize stover), wilting was actually found to give higher live-weight gain and feed efficiency. Conversely, with a high quality basal diet (Napier grass) wilting enhanced intake but not live-weight gain, so feed efficiency was greater for fresh material. The difference between fresh and wilted leaves was not great enough to justify the current widespread recommendation that calliandra should always be fed fresh.

The effects of seed mix and management on the abundance of desirable and pernicious unsown species in arable buffer strip communities

Buffer strips are refuges for a variety of plants providing resources, such as pollen, nectar and seeds, for higher trophic levels, including invertebrates, mammals and birds. Margins can also harbour plant species that are potentially injurious to the adjacent arable crop (undesirable species). Sowing perennial species in non-cropped buffer strips can reduce weed incidence, but limits the abundance of annuals with the potential to support wider biodiversity (desirable species). We investigated the responses of unsown plant species present in buffer strips established with three different seed mixes managed annually with three contrasting management regimes (cutting, sward scarification and selective graminicide). Sward scarification had the strongest influence on the unsown desirable (e.g. Sonchus spp.) and unsown pernicious (e.g. Elytrigia repens) species, and was generally associated with higher cover values of these species. However, abundances of several desirable weed species, in particular Poa annua, were not promoted by scarification. The treatments of cutting and graminicide tended to have negative impacts on the unsown species, except for Cirsium vulgare, which increased with graminicide application. Differences in unsown species cover between seed mixes were minimal, although the grass-only mix was more susceptible to establishment by C. vulgare and Galium aparine than the two grass and forb mixes. Annual scarification can enable desirable annuals and sown perennials to co-exist, however, this practice can also promote pernicious species, and so is unlikely to be widely adopted as a management tool in its current form.

Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins

P>1. Management of lowland mesotrophic grasslands in north-west Europe often makes use of inorganic fertilizers, high stocking densities and silage-based forage systems to maximize productivity. The impact of these practices has resulted in a simplification of the plant community combined with wide-scale declines in the species richness of grassland invertebrates. We aim to identify how field margin management can be used to promote invertebrate diversity across a suite of functionally diverse taxa (beetles, planthoppers, true bugs, butterflies, bumblebees and spiders). 2. Using an information theoretic approach we identify the impacts of management (cattle grazing, cutting and inorganic fertilizer) and plant community composition (forb species richness, grass species richness and sward architecture) on invertebrate species richness and body size. As many of these management practices are common to grassland systems throughout the world, understanding invertebrate responses to them is important for the maintenance of biodiversity. 3. Sward architecture was identified as the primary factor promoting increased species richness of both predatory and phytophagous trophic levels, as well as being positively correlated with mean body size. In all cases phytophagous invertebrate species richness was positively correlated with measures of plant species richness. 4. The direct effects of management practices appear to be comparatively weak, suggesting that their impacts are indirect and mediated though the continuous measures of plant community structure, such as sward architecture or plant species richness. 5. Synthesis and applications. By partitioning field margins from the remainder of the field, economically viable intensive grassland management can be combined with extensive management aimed at promoting native biodiversity. The absence of inorganic fertilizer, combined with a reduction in the intensity of both cutting and grazing regimes, promotes floral species richness and sward architectural complexity. By increasing sward architecture the total biomass of invertebrates also increased (by c. 60% across the range of sward architectural measures seen in this study), increasing food available for higher trophic levels, such as birds and mammals.

Numerical abundance of invasive ants and monopolisation of exudate-producing resources - a chicken and egg situation

1. Invasive ants commonly reach abnormally high abundances and have severe impacts on the ecosystems they invade. Current invasion theory recognises that not only negative interactions, such as natural enemy release, but positive interactions, such as facilitation, are important in causing this increased abundance. 2. For invasive ants, facilitation can occur through mutualism with exudate-producing plants and insects. To obtain such partnerships, however, invaders must first displace native ants, whose communities are highly structured around such resources. 3. By manipulating the abundance of an invasive ant relative to a native, we show that a minimum threshold abundance exists for invasive ants to monopolise exudate-producing resources. In addition, we show that behavioural dominance is context dependent and varies with spatial location and numerical abundance. 4. Thus, we suggest a 'facilitation-threshold' hypothesis of ant invasion, whereby a minimum abundance of invasive ants is required before facilitation and behavioural dominance can drive abundance rapidly upwards through positive feedback.