The effects of habitat structure on predation risk of birds in agricultural landscapes

It has been suggested that increased predation rates may rival habitat alteration as a causal agent in farmland bird population declines. Such a view may be over-simplistic, however, as changes in habitat structure may influence habitat selection and foraging efficiency through their influence on perceived and actual predation risk. We review evidence from the literature on the effects of habitat structure on predation risk of foraging and nesting birds and apply these principles to investigate the likely effects on the 20 species that comprise the UK Government's 'Farmland Bird Index'. Shorter vegetation is likely to enhance foraging efficiency and reduce predation risk (when ground foraging) for 15 of the 20 species. However, within grassland systems longer vegetation is known to enhance food supplies (e.g. Tipulid larvae and voles) of several farmland bird species and so mosaics of short and long vegetation may provide the optimum conditions for most species (e.g. Lapwing Vanellus vanellus , Starling Sturnus vulgaris , Barn Owl Tyto alba ). Agricultural intensification has encouraged uniform dense swards, thus reducing habitat diversity, and agri-environment schemes that provide heterogeneous sward structure may thus facilitate farmland bird conservation. Intensification has also resulted in less dense hedgerows; although a reversal of this trend may improve foraging efficiency for many species, it may be detrimental to a smaller number of species that prefer shorter, less dense hedges for nesting. Before these tentative conclusions can be confirmed, more research is required that considers how the effects of habitat structure on individuals is likely to translate into population-level impacts.     

Habitat use by seed-eating birds: a scale-dependent approach

The seedbank in arable farmland represents an important foraging resource for birds, particularly in grassland landscapes where alternative foraging opportunities may be scarce. We used a stepwise approach to examine the importance of seed food resources for farmland birds in winter. First, results from a large-scale experiment in which seed resources were manipulated to test the notion that birds aggregate at food resources, subject to a minimum threshold level. Secondly, a multiscale approach was used to characterize habitat use at a landscape scale and how this may inform agri-environment implementation. Overall seed resources declined sharply over the winter and were relatively low in most arable fields. Large-scale declines in the arable seedbank mean that much habitat may not be of sufficient quality to support foraging bird populations through a winter. At a landscape scale, extensive analyses of breeding season abundance show that bird abundance is most influenced by arable (i.e. seed-rich) habitat in grassland landscapes. The scale at which birds respond to habitat differs between species, and is related to the extent of between-season movements. Implementation of agri-environment schemes will need to consider both the quality of habitat and the context in which it is provided if recent declines in farmland bird populations are to be reversed.     

The Origin of Winter Visitors to the British Isles 7. Fieldfare (Turdus pilaris)

Capsule Large‐scale intensification of agricultural management during the past 50 years has resulted in a reduction of invertebrate abundance and higher and denser ground vegetation. Food availability for insectivorous birds foraging on the ground has been negatively affected, but the interactions between birds and their food availability are complex and often species‐specific. Populations of Wrynecks Jynx torquilla are declining all over Europe, possibly because of reduced accessibility to their main prey, ground‐dwelling ants, due to higher and denser ground vegetation. However, it is not clear which ground vegetation structures are tolerated by foraging Wrynecks and which habitats are preferred.

Aims To identify the optimal ground vegetation structure and the main habitat types in which Wrynecks search for food.

Method We radiotracked seven Wrynecks in high‐intensity farmland in Switzerland to study foraging habitat use during the reproduction season. Several habitat variables were mapped at each foraging location and compared with locations selected randomly within individual home ranges.

Results Wrynecks preferentially foraged at places with ≥50% bare ground. Vegetation height was not important. Older fruit tree plantations and fallow land were the preferred foraging habitats.

Conclusion Conservation measures should concentrate on preserving semi‐open agricultural landscape matrices with loose ground vegetation cover to provide suitable foraging conditions. This can be achieved even in intensively managed farmland as illustrated in this study.     

Habitat preferences of the Ortolan Bunting (Emberiza hortulana) in its prime wintering grounds,the cereal-dominated Ethiopian Highlands

Agricultural intensification and land-use changes are major factors impacting farmland biodiversity. The Ortolan Bunting Emberiza hortulana is the long-distance trans-Saharan migratory passerine that has undergone the most dramatic decline among all European farmland birds. The factors responsible for this decline may originate from the breeding grounds, migration stopovers and/or overwintering quarters. Very little is known about conditions on the species' wintering grounds, but a recent study has highlighted the utmost importance of the traditionally managed agroecosystems in the Ethiopian Highlands as a key wintering area, apparently harbouring as much as 90% of the world's Ortolan Bunting population. Using radiotracking and line transect surveys, this study aimed to provide fine-grained information about species–habitat relationships in the Ortolan Bunting overwintering quarters. Our results showed the importance, at the landscape scale, of small-scale agriculture, notably of traditionally managed, cereal-dominated fields interspersed with semi-natural structures. At a foraging-site scale, on the other hand, patches of bare ground in combination with large areas of post-harvesting stubble represented key habitat features. Stubbles provide an essential food resource and bare ground promotes ground foraging by enhancing food accessibility. The maintenance of a traditional agricultural economy will be essential to maintain the habitat potential for the Ortolan Buntings overwintering in the Ethiopian Highlands and will be instrumental in preserving its world population from further decline.     

Habitat selection by Ortolan Buntings Emberiza hortulana in post-fire succession in Catalonia: implications for the conservation of farmland populations

The Ortolan Bunting Emberiza hortulana is a long-distance migrant that has suffered major population declines across much of its European breeding range. While northern populations are bound largely to farmland, Mediterranean populations are largely confined to habitats subject to recurrent wildfires. Habitat selection of the Ortolan Bunting was assessed in a recently burnt area in Catalonia at landscape and habitat scales. A Zero-inflated Poisson procedure was used to model the abundance of birds in relation to landscape and habitat variables. The most parsimonious landscape model predicted the highest abundance on south-facing slopes, with a gradient above 10°. The most parsimonious habitat model showed a positive quadratic effect of bare ground and regenerating oak Quercus spp., with predicted optima for abundance around 20–30% and 20% cover, respectively. There was a clear relationship between predicted abundance of the Ortolan Bunting and post-fire regenerating oak shrubs. South-facing, moderately sloping areas were favoured and bare ground was a key feature of the species' habitat. A matrix combining patches of sparse oak shrubs and patches of bare ground appears to be the optimal breeding habitat in the Mediterranean. The maintenance or provision of similar habitat features, especially patches of bare ground, may prove crucial for the conservation of rapidly declining Ortolan Bunting populations on farmland across temperate Europe.     

Mixed biodiversity benefits of agri-environment schemes in five European countries

Agri-environment schemes are an increasingly important tool for the maintenance and restoration of farmland biodiversity in Europe but their ecological effects are poorly known. Scheme design is partly based on non-ecological considerations and poses important restrictions on evaluation studies. We describe a robust approach to evaluate agri-environment schemes and use it to evaluate the biodiversity effects of agri-environment schemes in five European countries. We compared species density of vascular plants, birds, bees, grasshoppers and crickets, and spiders on 202 paired fields, one with an agri-environment scheme, the other conventionally managed. In all countries, agri-environment schemes had marginal to moderately positive effects on biodiversity. However, uncommon species benefited in only two of five countries and species listed in Red Data Books rarely benefited from agri-environment schemes. Scheme objectives may need to differentiate between biodiversity of common species that can be enhanced with relatively simple modifications in farming practices and diversity or abundance of endangered species which require more elaborate conservation measures.     

Farmers’ perceptions of biodiversity and their willingness to enhance it through agri-environment schemes: A comparative study from Estonia and Finland

This comparative study is based on structured interviews with farmers participating in the agri-environment schemes in Estonia and Finland. It explores farmers’ interest in and knowledge of farmland wildlife, their understanding of the concept of biodiversity, and awareness of the potential causes behind declines of farmland birds. It also examines the relationship between farmers’ interest and willingness to undertake practices favouring farmland wildlife. Estonian and Finnish farmers showed considerable interest in wildlife on their farms, which was only weakly related to the self-assessed knowledge of wildlife. Many farmers viewed biodiversity from a narrow perspective often excluding species directly related to farming. Finnish farmers expressed more concern about the decline in common farmland species than Estonian ones. In both countries farmers rated intensification of agriculture as the major driving force behind farmland bird declines. The expressed interest in wildlife positively correlated with willingness to undertake wildlife-friendly measures. Only farmers with agri-environment contracts targeted specifically at biodiversity enhancement were more knowledgeable about practical on-farm activities favouring wildlife, and were more willing to employ them than the rest. Estonian farmers expressed a high degree of willingness to enhance wildlife though agri-environment management, which is a good sign for better implementation of the recently established national programme towards conservation.     

Winter availability of cereal stubbles attracts declining farmland birds and positively influences breeding population trends

Many studies have demonstrated the selection of stubble fields by farmland birds in winter, but none have shown whether provisioning of this key habitat positively influences national population trends for widespread farmland birds. We use two complementary extensive bird surveys undertaken at the same localities in summer and winter and show that the area of stubble in winter attracts increased numbers of several bird species of conservation concern. Moreover, for several farmland specialists, the availability of stubble fields in winter positively influenced the 10 year breeding population trend (1994-2003) whereas hedgerow bird species were less affected. For skylarks and yellowhammers, initially negative trends showed recovery with 10-20 ha of stubble per 1 km square. Thus, agri-environment schemes that promote retention of over-winter stubbles will attract birds locally and are capable of reversing current population declines if stubbles are available in sufficient quantity.     

The management of crop structure: a general approach to reversing the impacts of agricultural intensification on birds?

Agricultural intensification is accepted widely as a cause of bird population declines on farmland in Europe and North America. Although intensification is multivariate, one common theme is the impact on variation in crop structure, both within and between fields. Intensification creates simpler, more homogeneous and denser swards in both tillage crops and grassland. This influences predation risk, exposure to weather extremes and the diversity, abundance and accessibility of food. Birds trade off these pressures in different ways, so that the more uniform and dense the vegetation, the fewer the number of birds and range of species that are able to nest and forage successfully. Reversing recent trends towards dense, simplified and homogeneous swards will improve nesting and foraging habitat conditions for a wide range of species across farming systems, and may represent a cost-effective mechanism for the further improvement of agri-environment scheme options designed to assist the recovery of farmland bird populations.     

The relevance of non-farmland habitats, uncropped areas and habitat diversity to the conservation of farmland birds

Use of non-farmland habitats by species generally perceived as 'farmland birds' is common, yet these habitats are not always considered in conservation strategies aimed at population recovery. At the national scale, many farmland species occur in landscapes not dominated by farmland. An analysis of distribution atlas data coupled with remotely sensed habitat data showed that for 16 out of 28 farmland species, less than half of the breeding range was associated with high cover of lowland farmland. However, with a few exceptions, populations breeding in non-farmland habitats are likely to depend on farmland at some time in the year. Within farmland landscapes, uncropped areas and patches of non-farmland habitat can provide nesting, foraging or roosting resources. Habitats that are scarce on farmland and that provide potential supplementary or complementary resources to those available within the productive areas of farmland include ruderal vegetation, rough grassland and scrub. Enhancing habitat diversity through provision of modest quantities of these habitats will benefit farmland birds. Complete knowledge of year-round habitat requirements and patterns of resource use at all scales is needed if robust national conservation plans are to be developed for farmland species. Similarly, interactions between the farmland and non-farmland sections of populations need to be determined.     

Fine-scale habitat structure complexity determines insectivorous bird diversity in a tropical forest

Habitat complexity in reforested stands has been acknowledged as a key factor that influences habitat use by birds, being especially critical for habitat disturbance-sensitive species such as tropical understory insectivorous birds. Most studies regarding the relationship between forest structure and species diversity were conducted at the landscape scale, but different diversity patterns may emerge at a finer scale (i.e., within a habitat patch). We examined a tropical reforested area (State of Caldas, Colombia), hypothesizing that insectivorous bird richness, abundance, and foraging guild abundance would increase as intra-habitat complexity increases. We established 40 monitoring plots within a reforested area, measured their structural features, and determined their relationships with species richness, total abundance, and foraging guild abundance, using Generalized Additive Models. We found that the increasing variation in basal area, stem diameter, and number of stems was positively correlated with species richness, total abundance, and foraging guild abundance. Relationships between richness or abundance and structural features were not lineal, but showing curvilinear responses and thresholds. Our results show that heterogeneity on basal area, stem diameter, and the number of stems was more correlated to insectivorous bird richness and abundance than the average of those structural features. Promoting structural variation on reforested areas by planting species with different growth rates may contribute to increase the richness and abundance of a tropical vulnerable group of species such as the understory insectivorous birds.     

The consequences of predator control for brown hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) on UK farmland

The brown hare Lepus europaeus is a valued game species but also a species of conservation concern owing to its severe decline in abundance on farmland throughout Europe during the twentieth century. Changes in the farmland habitat and predation have both been cited as causative factors. Their relative roles have been unclear, but most conservation action has focused on improving habitat. We analyse data from a sequence of three unique studies (one experiment and two demonstrations) covering the period 1985–2006 in which control of several common predator species was undertaken to increase densities of wild game on farmland in England. Across the three studies, regression modelling of the proportional change in hare numbers between successive years showed that—after site, year differences and harvesting were accounted for—predator control was a significant determinant of hare population change. Where habitat improvement also took place, hares reached autumn densities that were exceptional for the UK and which could sustain substantial harvests. When predation control was stopped, hare densities fell, even where habitat improvements remained in place. This analysis demonstrates that even where farmland habitat is greatly improved, uncontrolled predation prevents hares making full use of its carrying capacity. This helps explain the mixed—and at best modest—success of agri-environment schemes in the UK and elsewhere in Europe to increase hare densities. Game-shooting estates, on which effective predator control takes place, probably have a special significance within the landscape as source areas for brown hares.     

Insectivorous birds are not effective pest control agents in olive groves

Although avian-mediated pest control is a significant ecosystem service with important economic implications, few experimental studies have ever documented its role in Mediterranean agroforests. Specifically, information on pest control by birds is lacking in certain permanent agroecosystems of worldwide importance such as olive groves.Here, we assess experimentally for the first time the effectiveness of insectivorous birds in controlling the two main olive-tree pests. We also explore the effects of distance to semi-natural habitat patches on avian insectivore abundance and pest control. We combined bird and pest surveys with pest damage monitoring and two field experiments (branch exclusion and plasticine models) at a regional scale.The experiments showed that birds played a negligible role as pest controllers (measured in terms of attack rates on plasticine models and controlled pest damage) in the studied olive groves; overall, pests were abundant and pest damage was high on most farms. In addition, surveys showed that insectivorous birds were more abundant and diverse in patches of semi-natural habitat, compared to the matrix of olive groves, and that proximity to semi-natural patches was not a driver of bird-driven pest control.This study experimentally demonstrates that insectivorous birds are not effective pest controllers in olive groves. The absence of patterns linking insectivorous birds’ availability and observed pest control suggests that birds are unable to exert effective control over the main olive-tree pests. This lack of biocontrol by birds is probably due to low accessibility and/or appetence for the current insectivorous groups. Habitat improvement aimed at encouraging some under-represented forager species could improve the likelihood that birds will provide this ecosystem service.     

The effect of habitat complexity on the functional response of a seed-eating passerine

Recent population declines of seed-eating farmland birds have been associated with reduced overwinter survival due to reductions in food supply. An important component of predicting how food shortages will affect animal populations is to measure the functional response, i.e. the relationship between food density and feeding rate, over the range of environmental conditions experienced by foraging animals. Crop stubble fields are an important foraging habitat for many species of seed-eating farmland bird. However, some important questions remain regarding farmland bird foraging behaviour in this habitat, and in particular the effect of stubble on farmland bird functional responses is unknown. We measured the functional responses of a seed-eating passerine, the Chaffinch Fringilla coelebs , consuming seeds placed on the substrate surface in three different treatments: bare soil, low density stubble and high density stubble. Stubble presence significantly reduced feeding rates, but there was no significant difference between the two stubble treatments. Stubble reduced feeding rates by reducing the maximum attack distance, i.e. the distance over which an individual food item is targeted and consumed. The searching speed, handling time per seed, proportion of time spent vigilant, duration of vigilance bouts and duration of head-down search periods were unaffected by the presence of stubble. The frequency of vigilance bouts was higher in the bare soil treatment, but this is likely to be a consequence of the increased feeding rate. We show the influence of a key habitat type on the functional response of a seed-eating passerine, and discuss the consequences of this for farmland bird conservation.     

Is breeding of farmland wading birds depressed by a combination of predator abundance and grazing?

Agri-environment schemes have been implemented across Europe to counter biodiversity loss in agricultural landscapes and halt the continual decline of farmland birds, including waders. Such schemes provide financial compensation for changes in agricultural practice, including livestock grazing regimes. Scheme uptake has been variable, partly because farmers believe that other factors, notably predation, are key to wader population declines. On the basis of wader breeding surveys across Shetland, UK, we show that predator density and livestock grazing, through reducing sward height, interact to influence territoriality and thereby are likely to affect wader breeding success. Our results appear to reflect views of both farmers and government agencies, which indicates that future agri-environment schemes would benefit from genuine stakeholder participation to maximize scheme uptake, implementation and beneficial effects on biodiversity. Our findings also imply that agri-environment schemes will reap the greatest benefits for waders through reducing stocking rate where avian predators are abundant.     

The role of farm structure on bird assemblages around a Kenyan tropical rainforest

Although it is clear that the farmlands neighbouring fragmented forests are utilized by some forest birds, it is not clear how birds in general respond to farmland habitat mosaic. An effort was made to determine how bird density and foraging assemblages were influenced by farm structural characteristics and distance from forest edge. Thirty farms up to a distance of 12 km around Kakamega forest in western Kenya were studied. Farm structure entailed size, hedge volume, habitat heterogeneity, woody plant density, plant diversity and crop cover. Birds were surveyed using line transects and DISTANCE analyses and classified into six feeding guilds and three habitat associations. Size of farms increased away from the forest, as woody plant density, plant diversity, indigenous trees and subsistence crop cover declined. The most important farm structure variable was hedge volume, which enhanced bird species richness, richness of shrub‐land bird species and insectivorous bird density (R = 0.58, P < 0.01). Bird density increased with tree density while indigenous trees were suitable for insectivores and nectarivores. There were very few forest bird encounters. Agricultural practices incorporating maintenance of hedges and sound selection of agroforestry trees can enhance conservation of birds on farmland, though, not significantly for forest species.     

Birds in willow short-rotation coppice compared to other arable crops in central England and a review of bird census data from energy crops in the UK

We present the results of spring and winter surveys of birds using 22 willow short-rotation coppice (SRC) fields and 22 paired arable or grassland controls over a 5 year period in central and northern England. The study sites were linked to the first commercial wood-fuelled electricity plant in Yorkshire and were managed commercially. We use the findings of this study, in the context of other work on birds in energy crops, to suggest how birds may be affected by commercial cropping of SRC or energy grasses on UK farmland. For willow SRC, most existing farmland species were not completely displaced by SRC cropping. Even farmland specialists such as Skylark Alauda arvensis used cut SRC fields. Some species characteristic of scrub and wood-type habitats were attracted onto farmland by SRC. Birds were more abundant in hedgerows next to SRC than in those adjoining arable or grass. The interior of large SRC plots contained fewer birds than the edge-zone (< 50 m). In winter, the same resident species occured in SRC as the summer, plus winter migrants such as Snipe Gallinago gallinago and Fieldfare Turdus pilaris . Overall, more individuals and species were recorded in and around SRC than equivalent arable or grassland throughout the year. Yellow Wagtail Motacilla flava and certain locally occurring farmland specialists may be displaced by SRC plantations. Equivalent census data for energy grasses in the UK are not available. The impact of large regional energy crop developments on birds is not known. We do not know if species commonly recorded in these crops survive and breed well so population effects are unknown. However SRC crops are often weedy and insect rich and hence have potential as foraging habitats in summer and winter. The scope for agri-environment schemes to include energy crops is discussed.     

Dung‐associated arthropods influence foraging ecology and habitat selection in Black‐necked Cranes (Grus nigricollis) on the Qinghai–Tibet Plateau

Variation in grassland vegetation structure influences the habitat selection of insectivorous birds. This variation presents a trade‐off for insectivorous predators: Arthropod abundance increases with vegetation height and heterogeneity, but access to arthropod prey items decreases. In contrast, grazing by large herbivores reduces and homogenizes vegetation, decreasing total arthropod abundance and diversity. However, the presence of livestock dung may help counteract the overall reduction in invertebrates by increasing arthropods associated with dung. It is unclear, however, how the presence of arthropod prey in dung contributes to overall habitat selection for insectivorous birds or how dung‐associated arthropods affect trade‐offs between vegetation structure, arthropod abundance, and access to prey. To explore these relationships, we studied habitat selection of the Black‐necked Crane (Grus nigricollis), a large omnivorous bird that breeds on the Qinghai–Tibet Plateau. We assessed the relationships between habitat selection of cranes and vegetation structure, arthropod abundance, and the presence of yak dung. We found that Black‐necked Cranes disproportionately foraged in grassland patches with short sward height, low sward height heterogeneity, and high numbers of dry yak dung, despite these habitats having lower total arthropod abundance. Although total arthropod abundance is lower, these habitats are supplemented with dry yak dung, which are associated with coleopteran larvae, making dung pats an indicator of food resources for breeding Black‐necked Cranes. Coleopteran adults and larvae in yak dung appear to be an important factor influencing the habitat selection of Black‐necked Cranes and should be considered when assessing grassland foraging trade‐offs of insectivorous birds. This research provides new insights into the role of livestock dung in defining foraging habitats and resources for insectivorous predators.     

Functional Invertebrate Prey Groups Reflect Dietary Responses to Phenology and Farming Activity and Pest Control Services in Three Sympatric Species of Aerially Foraging Insectivorous Birds

Farming activity severely impacts the invertebrate food resources of farmland birds, with direct mortality to populations of above-ground arthropods thorough mechanical damage during crop harvests. In this study we assessed the effects of phenological periods, including the timing of harvest, on the composition and biomass of prey consumed by three species of aerial insectivorous birds. Common Swifts Apus apus, Barn Swallows Hirundo rustica and House Martins Delichon urbica breed sympatrically and most of their diet is obtained from agricultural sources of invertebrate prey, especially from oil-seed rape crops. We categorized invertebrate prey into six functional groups, including oil-seed rape pests; pests of other arable crops; other crop-provisioned taxa; coprophilous taxa; and taxa living in non-crop and mixed crop/non-crop habitats. Seasonality impacted functional groups differently, but the general direction of change (increase/decrease) of all groups was consistent as indexed by prey composition of the three aerial insectivores studied here. After the oil-seed rape crop harvest (mid July), all three species exhibited a dietary shift from oil-seed rape insect pests to other aerial invertebrate prey groups. However, Common Switfts also consumed a relative large quantity of oil-seed rape insect pests in the late summer (August), suggesting that they could reduce pest insect emigration beyond the host plant/crop. Since these aerially foraging insectivorous birds operate in specific conditions and feed on specific pest resources unavailable to foliage/ground foraging avian predators, our results suggest that in some crops like oil-seed rape cultivations, the potential integration of the insectivory of aerial foraging birds into pest management schemes might provide economic benefits. We advise further research into the origin of airborne insects and the role of aerial insectivores as agents of the biological control of crop insect pests, especially the determination of depredation rates and the cascading effects of insectivory on crop damage and yield.