Assessing breeding success in common woodland birds using a novel method

Capsule Repeated counts of fledged broods can provide a useful estimate of breeding success for most common woodland birds.

Aims To assess the efficacy of comparing fledged-brood survey data with territory mapping using simple mark–recapture analysis techniques to provide an estimate of breeding success for common woodland birds that does not involve finding nests.

Methods Three observers undertook territory mapping surveys of adults, followed by counts of fledged broods four times a week during May–July 2007 in two 15 ha woods each, both in southern England. Using known fledging to maturity periods, these counts were used to calculate daily detection probabilities for broods of ubiquitous species. These enabled fledged brood territory occupancy probabilities (i.e. brood to territory ratios) to be estimated that take account of the possibility that broods were present but missed by surveys.

Results Of the 19 species found in all six woods, mean daily detection probability estimates for fledged broods of 17 species ranged from 0.17 to 0.50 with significant variation between woods for 12 species, but within region/observer for four species. The mean probability of detecting a brood at least once was over 75% using four visits per week and over 50% using two visits. Only for Great Spotted Woodpeckers Dendrocopos major and Garden Warblers Sylvia borin was the fledging period too short and the daily detection probability too low to provide a reasonable estimate of the territory occupancy probability.

Conclusion Daily detection probabilities for fledged broods of most common woodland birds were sufficiently high to enable useable estimates of fledged-brood territory occupancy probabilities to be made based on a survey programme involving two or three visits per week between late May and the end June. The method used may have application as a means of providing a relatively easily derived productivity index for woodland bird monitoring programmes or for research studies.     

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 optimal age of sown field margins for breeding farmland birds

The Europe‐wide decline in the populations and diversity of farmland birds has not been stopped despite dedicated conservation efforts such as agri‐environment schemes (AES). The main reason for the lack of success of AES is considered to be their low ecological quality and insufficient area. Understanding the effects of different management strategies on the ecological quality of AES is therefore important. Here, we investigate the relationship between breeding bird density and species richness and the age of sown field margins, a widely used type of AES, in southwestern Switzerland. Territories of breeding birds were mapped on 67 field margins between 2004 and 2011. Territory densities (for eight species) and species richness were analysed in relation to age of the field margin. A general negative correlation between size of the field margin and territory density indicated that territory density was higher when the birds could forage in adjacent cultivated land. Territory densities and species richness increased up to an age of 4–6 years after sowing, depending on the species, and declined thereafter. The results suggest that the co‐occurrence of newly sown margins and margins over 3 years old will have a positive effect on breeding bird densities and species diversity.     

Within-season sequential density dependence regulates breeding success in mallards Anas platyrhynchos

Density dependence in vital rates is a key issue in population ecology but remains largely unexplored experimentally. We studied breeding success, lake use, and prey availability in wild mallards Anas platyrhynchos on small nemoral lakes in a replicated, two-year cross-over experiment in which pair density was increased. The number of wild mallards that settled on lakes prior to introductions of extra pairs did not differ between control and introduction years. Introductions led to a lake-level reduction in the number of broods observed. However, the number of stage 2+ (almost fledged) ducklings did not differ between treatments, nor did lake utilization by nonbreeding adults, broods and ducklings. Prey resource availability differed greatly among lakes, but it did not correlate with breeding success. Partialling out the possible effect of food competition from wild adult nonbreeding mallards did not change this conclusion. Our study demonstrates sequential density dependence in breeding success; introductions caused a decrease in brood number, but despite fewer broods a similar number of nearly fledged ducklings were produced. We suggest that predation and/or lake change of broods soon after hatching created these patterns. We conclude that using a single and late measure of breeding success such as fledged birds can mask regulatory processes. Implications of density dependence and its relation to individual reproductive success are understood better if breeding success is decomposed into nest success, duckling survival and fledgling survival.     

Effects of perennial wildflower strips and landscape structure on birds in intensively farmed agricultural landscapes

Farmland bird populations are in a deep crisis across Europe. Agri-environment schemes (AES) were implemented by the European Union to stop and reverse the general decline of biodiversity in agricultural landscapes. In Germany, flower strips are one of the most common AES. Establishing high-quality perennial wildflower strips (WFS) with species-rich native forb mixtures from regional seed propagation is a recent approach, for which the effectiveness for birds has not yet been sufficiently studied. We surveyed breeding birds and vegetation on 40 arable fields with WFS (20 with single and 20 with aggregated WFS) and 20 arable fields lacking WFS as controls across Saxony-Anhalt (Germany). Additionally, vegetation composition, WFS quantity and landscape structure (e.g. distance to nearest woody element) were considered in our analyses. All WFS were established with species-rich native seed mixtures (30 forbs) in agricultural practice as AES. Arable fields with WFS had a higher species richness and territory density of birds than controls, confirming the effectiveness of this AES. A forb-rich vegetation was the main driver promoting birds. Flower strip quantity at the landscape level had positive effects only on bird densities, but also single WFS achieved benefits. A short distance from WFS to woody elements increased total bird species richness. However, the density of farmland birds, which are target species of these AES, were negatively affected by the proximity and proportion of woody elements in the vicinity. The effect of the proportion of non-intensively used open habitats and overall habitat richness was unexpectedly low in the otherwise intensively farmed landscape. Species-rich perennial WFS significantly promoted breeding birds. Successful establishment of WFS, resulting in high-quality habitats, a high flower strip quantity as well as implementation in open landscapes were shown to maximise the effectiveness for restoring declining and AES target farmland birds.     

The Impact of Increased Food Availability on Reproduction in a Long-Distance Migratory Songbird: Implications for Environmental Change?

Many populations of migratory songbirds are declining or shifting in distribution. This is likely due to environmental changes that alter factors such as food availability that may have an impact on survival and/or breeding success. We tested the impact of experimentally supplemented food on the breeding success over three years of northern wheatears (Oenanthe oenanthe), a species in decline over much of Europe. The number of offspring fledged over the season was higher for food-supplemented birds than for control birds. The mechanisms for this effect were that food supplementation advanced breeding date, which, together with increased resources, allowed further breeding attempts. While food supplementation did not increase the clutch size, hatching success or number of chicks fledged per breeding attempt, it did increase chick size in one year of the study. The increased breeding success was greater for males than females; males could attempt to rear simultaneous broods with multiple females as well as attempting second broods, whereas females could only increase their breeding effort via second broods. Multiple brooding is rare in the study population, but this study demonstrates the potential for changes in food availability to affect wheatear breeding productivity, primarily via phenotypic flexibility in the number of breeding attempts. Our results have implications for our understanding of how wheatears may respond to natural changes in food availability due to climate changes or changes in habitat management.     

Effects of changes in farming of subalpine meadows on a grassland bird, the whinchat ( Saxicola rubetra)

The increasing intensity of farming of meadows is supposed to be the major cause for strong declines of many meadow breeding birds. The whinchat, Saxicola rubetra, a Palaearctic migratory bird, is an indicator species of open grassland farmed at a low intensity. Originally widespread throughout Switzerland, it is now restricted to mountain and subalpine grassland. We document the changes in meadow cultivation in subalpine farmland from 1988 to 2002, and the breeding performance and density of the whinchat. We explored the impact of habitat degradation on the population dynamics of this meadow bird. The cultivation of hay meadows changed markedly within the 15 years: the onset of mowing was shifted forward by about 20 days, and farmers applied new techniques such as silage and irrigation. This shift was more pronounced in the favourable farmland in the valley bottom (Pradellas) than on the slopes (Vnà). The percentage of successful whinchat broods, ranging from 5% to 78% in different years, strongly depended on mowing date on both sites. In spite of earlier mowing, birds did not change their time schedule of breeding. Breeding success in Pradellas was too low to compensate for mortality (sink population), but because of immigration the number of breeding pairs did not decrease untill 2000. Population size may therefore be a misleading indicator of local population viability. Based on the breeding schedule of whinchats at different altitudes, we recommend mowing dates in agreement with the reproductive cycle of ground nesting meadow birds.     

Habitat preference and mobility of <Emphasis Type="Italic">Polia bombycina</Emphasis>: are non-tailored agri-environment schemes any good for a rare and localised species?

General agri-environment schemes (AES) have been shown to benefit widespread species, but there is little information on the extent to which rare, more localised, species may also benefit. We tested whether AES options aimed at increasing general biodiversity also benefit a highly endangered moth, Polia bombycina, without species-specific tailoring. We assessed effects on its abundance of two AES options, wide field margins and hedgerow trees, using light traps at the landscape-scale and for mark-release-recapture at the farm-scale. We hypothesized that abundance would be highest at wide field margins and at hedgerow trees, and that if hedgerow trees conferred a positive effect, individuals would be more likely to follow hedgerows than crossing exposed fields while on the move. The results showed that significantly more individuals were captured at sites with a hedgerow tree. Numbers were also higher at wide margins, but this was not statistically significant, and no individuals were caught at field centres. Our study suggests that general options within appropriately designed and implemented AES aimed at increasing overall biodiversity in intensive agricultural landscapes have the potential to not only benefit common, widespread habitat generalists, but some rare and more endangered species as well. P. bombycina serves as an example of how general AES options, existing and novel ones alike, might cater for the needs of rare and localised species. As the precise ecological requirements of most invertebrate species remain unknown, we urge scientists and governments to address the challenge to research and design truly general AES, which options should be able to deliver not only for widespread species but also for the less-widespread counterpart of farmland biodiversity.     

Should conservation strategies consider spatial generality? Farmland birds show regional not national patterns of habitat association

A key assumption underlying any management practice implemented to aid wildlife conservation is that it will have similar effects on target species across the range it is applied. However, this basic assumption is rarely tested. We show that predictors [nearly all associated with agri-environment scheme (AES) options known to affect European birds] had similar effects for 11 bird species on sites with differing farming practice (pastoral vs. mixed farming) or which differed in the density at which the species was found. However, predictors from sites in one geographical region tended to have different effects in other areas suggesting that AES options targeted at a regional scale are more likely to yield beneficial results for farmland birds than options applied uniformly in national schemes. Our study has broad implications for designing conservation strategies at an appropriate scale, which we discuss.     

Habitat type determines the effects of disturbance on the breeding productivity of the Dartford Warbler Sylvia undata

Numerous studies have examined the causes and impacts of human disturbance on birds, but little is known about how these impacts vary among habitats. This is of applied importance both for predicting bird responses to changes in disturbance and in planning how to reduce disturbance impacts. The Dartford Warbler Sylvia undata , a key heathland breeding species, occupies territories in a range of heathland types. Three territory habitat groups were identified: heather-dominated territories, heather territories with significant areas of European Gorse Ulex europaeus and territories containing Western Gorse U. gallii . Productivity was significantly affected by the timing of breeding in all habitats, but disturbance only appeared to have a significant impact on the productivity of birds in heather territories. Disturbance events in heather territories delayed breeding pairs for up to 6 weeks. This significantly decreased both the number of successful broods raised and the average number of chicks fledged per pair. Nests situated close to territory boundaries in heather territories, with high numbers of disturbance events, were more likely to fail outright. It was determined that an average of between 13 and 16 people passing through a heather territory each hour would delay breeding pairs sufficiently to prevent multiple broods.     

Assessing the impact of Entry Level Stewardship on lowland farmland birds in England

Agri‐environment schemes (AES) are central to the conservation of Europe’s farmland biodiversity. The UK Government’s Public Service Agreement target seeks to reverse the decline of farmland birds in England by 2020 through the use of AES. The Entry Level Stewardship (ELS) scheme, introduced in 2005, is the first non‐competitive, broad‐uptake stewardship scheme designed to deliver simple but effective environmental management on farms throughout England. Approximately 5 million hectares of farmland are currently entered into Environmental Stewardship (ES) agreements, and given the scale of this investment it is important to verify the efficacy of ELS as a mechanism for delivering biodiversity benefits. We used spatially referenced ES agreement data to assess the influence of specific management options and combinations of options on farmland bird population changes between 2005 and 2008 derived from standardized surveys carried out on over 2000 lowland farmland 1‐km squares in England. The results showed only limited evidence for short‐term effects of ES. Only Corn Bunting Emberiza calandra and Common Starling Sturnus vulgaris showed some landscape‐specific positive associations with area under ELS management. There was also limited evidence for positive associations between specific management options and the population changes of the species at which they were aimed. The current pattern of option uptake may be limiting benefits of ELS, with boundary management accounting for over 50% of scheme compensation value. However, the time required for option maturation and the time lags in bird population responses to environmental change mean that it is too early to reach definitive conclusions about the success or failure of the scheme. The findings of this study will assist in the development of ELS by feeding into the 5‐year review scheduled for 2010 and have implications for the development of similar schemes elsewhere in Europe.     

Moving Window Abundance – A method to characterise the abundance dynamics of farmland birds: The example of Skylark (Alauda arvensis)

Farmland birds are important indicators of the state of biodiversity in rural landscapes, and the occurrence and abundance of birds contribute to their importance as bioindicators. However, the measurement of farmland bird abundance can be difficult. The rapid growth of crop plants in the spring combined with disturbances related to farming practices, such as weed and pest control and other measures, can profoundly change the habitat suitability of arable fields for birds within a short period of time. Consequently, the existing dynamics must be incorporated into the applied methods, and a single value of bird abundance during the breeding season is insufficient to characterise the habitat functions of arable land. The abundance of farmland bird species is influenced by crop specific features, which profoundly change within the breeding season and which we have described using a novel concept called Moving Window Abundance. Based on field surveys in 29 observation areas within arable landscapes, each 1 km², Skylark with territorial behaviour were counted using a mapping method related to the habitats and growth of field crops, such as wheat and maize. To describe the dynamic characteristics of the abundance during the breeding season over the time period from 16 March to 18 July, three methods of Moving Window Abundance were tested: patchy, adjacent and overlapping. These methods differ in the time-space continuity of the considered time window throughout the breeding season. In the Patchy Moving Window Abundance method, we used the exact days in which field surveys were conducted throughout the entire time period of the field surveys. However, with the Adjacent Moving Window Abundance method, the time window included five-day, ten-day and semi-monthly adjacent windows; in the Overlapping Moving Window Abundance method, the time window shifted daily, with five-day, ten-day and semi-monthly overlapping windows used in the calculation procedure. The results indicated that the dynamic nature of Skylark abundance (i) reached a maximum level in the first breeding period in the agricultural landscape and (ii) exhibited large variations in level and time within the various field crops. Therefore, abundance was not described conventionally using a single numerical value but rather using mathematical functions based on the spatial scale of the landscape and habitat. We conclude with recommendations for further research to standardise farmland bird monitoring.     

Calcium shortage as a constraint on reproduction in great tits Parus major: a field experiment

Several bird species breeding in acidified habitats face serious problems caused by lowered availability of calcium-rich foods. Moreover, recent studies have revealed that calcium shortage causes phenotypic variation in certain reproductive parameters of passerines even in non-acidified habitats. However, to date, no clear evidence has been found of the limiting effect of calcium deficiency on the reproductive output of birds breeding in such habitats. In this paper, we present experimental evidence from a long-term study that calcium shortage may indeed limit reproductive output of passerines that breed in non-acidified habitats. We carried out calcium-supplementation experiments with great tits Parus major breeding in coniferous and deciduous woodlands in Estonia. Calcium-supplemented birds laid more eggs, tended larger broods and raised more fledglings with longer tarsi compared with control birds. Calcium shortage constrained reproduction of tits not only in spring, but also in mid-summer, during second breeding attempts. Moreover, no significant differences were detected between deciduous and coniferous habitats with respect to the effect of calcium supplements on reproductive success.     

Effects of agri-environmental schemes on farmland birds: do food availability measurements improve patterns obtained from simple habitat models?

Studies evaluating agri‐environmental schemes (AES) usually focus on responses of single species or functional groups. Analyses are generally based on simple habitat measurements but ignore food availability and other important factors. This can limit our understanding of the ultimate causes determining the reactions of birds to AES. We investigated these issues in detail and throughout the main seasons of a bird's annual cycle (mating, postfledging and wintering) in a dry cereal farmland in a Special Protection Area for farmland birds in central Spain. First, we modeled four bird response parameters (abundance, species richness, diversity and “Species of European Conservation Concern” [SPEC]‐score), using detailed food availability and vegetation structure measurements (food models). Second, we fitted new models, built using only substrate composition variables (habitat models). Whereas habitat models revealed that both, fields included and not included in the AES benefited birds, food models went a step further and included seed and arthropod biomass as important predictors, respectively, in winter and during the postfledging season. The validation process showed that food models were on average 13% better (up to 20% in some variables) in predicting bird responses. However, the cost of obtaining data for food models was five times higher than for habitat models. This novel approach highlighted the importance of food availability‐related causal processes involved in bird responses to AES, which remained undetected when using conventional substrate composition assessment models. Despite their higher costs, measurements of food availability add important details to interpret the reactions of the bird community to AES interventions and thus facilitate evaluating the real efficiency of AES programs.     

Effect of age,haemosporidian infection and body condition on pair composition and reproductive success in Great Tits Parus major

Several factors shape lifetime reproductive success, including genetic background, body condition, environmental conditions and ecological interactions such as parasitism. Adults often show higher reproductive success than their young conspecifics, especially in long-lived bird species, and this may be explained by the cumulative effects of an increase in reproductive experience and the selection of high-quality individuals from one year to the next. To test whether this pattern also exists in short-lived bird species, we used 13 years of monitoring data from two Great Tit Parus major populations. The effects of male and female age on several reproductive parameters were analysed in 419 pairs of Great Tits, while accounting for body condition and infection by haemosporidian parasites. Reproductive success was mainly affected by the age-class of males. Pairs containing a sub-adult male fledged one-third fewer chicks than pairs containing an adult male. The difference was not caused by variation in male fertility but could have been caused by better parental care provided by adult birds. In addition to lower reproductive success, first-year males also had reduced access to mating compared with adult males, suggesting an avoidance of sub-adult males by females. Nestling body condition was positively correlated with parental body condition, and the body condition of male and female members of breeding pairs was positively correlated. Finally, the number of fledged chicks was mainly affected by the infection status of males. These results temper our previously published results showing an effect of infection on Great Tit reproduction regardless of their sex. In our previous study, and as in most cases, the status of the partner was not taken into account and we show here that this is essential because it can lead to a biased interpretation of the results.     

Breeding biology and longevity of Russet-crowned Motmots in central Mexico

ABSTRACT.   Motmots, with their distinctive racketed-tails, represent one of the most easily recognized tropical birds, yet little is known about the basic natural history of most species in the family Momotidae. We report basic breeding biology and longevity of Russet-crowned Motmots ( Momotus mexicanus ), a medium-sized Neotropical bird that ranges from northwest Mexico to central Guatemala. We monitored nest success of eight pairs from 1 May to 17 July 1998 in tropical deciduous forests in central Mexico. Motmots laid an average of 4.1 eggs and incubated for approximately 20 d. Four of eight nests fledged young. Of these four nests, average hatching success was 69% and average fledgling success was 56%. The mean duration of the nestling period at three nests was 33.7 d. Based on the recapture of one individual bird in April 2008, we provide a longevity estimate that Russet-crowned Motmots can survive at least 11 yr in the wild. These data on nesting success and longevity add to our limited knowledge of the natural history of this understudied species.     

Modelling land-use sustainability using farmland birds as indicators

Biodiversity on farmland is declining due to agricultural intensification and occurs across many taxa such as plants, insects or birds. Here, we modelled population sizes of five farmland birds in central Germany as the German Sustainability Indicator for Species Diversity (SISD) is based on this taxon. We explored options for sustainable farmland management by generating land-use scenarios at the regional scale. For individual bird species, high SISD scores could be reached by changing environmental variables, such as landscape or crop diversity, percent cover of spring cereals or hedge density. However, contrasting species responses to these variables prevented from reaching high scores for all species simultaneously. We were able to improve the total SISD score from 0.77 at present to 0.94 by increasing landscape and crop diversity or to 0.87 by increasing hedge density and reducing spring cereals, respectively. An economic evaluation of the return losses associated with these changes revealed that annual costs of approx. 5.5 €/ha farmland would suffice for this latter increase by optimizing hedges and spring cereals towards high SISD. We conclude that balancing three levels of trade-offs, i.e. contrasting requirements of species, diverging responses in different landscapes, and alternative economic options, is difficult to achieve without systematic modelling. Yet, by accounting for differences in landscape structure and species distributions at a regional spatial scale and by focusing on clearly defined measures such as hedge density or the cover of spring cereals rather than composite indices like landscape or crop diversity it seems possible to develop realisable, affordable and sustainable management strategies.     

Anthropogenically-Mediated Density Dependence in a Declining Farmland Bird

Land management intrinsically influences the distribution of animals and can consequently alter the potential for density-dependent processes to act within populations. For declining species, high densities of breeding territories are typically considered to represent productive populations. However, as density-dependent effects of food limitation or predator pressure may occur (especially when species are dependent upon separate nesting and foraging habitats), high territory density may limit per-capita productivity. Here, we use a declining but widespread European farmland bird, the yellowhammer Emberiza citrinella L., as a model system to test whether higher territory densities result in lower fledging success, parental provisioning rates or nestling growth rates compared to lower densities. Organic landscapes held higher territory densities, but nests on organic farms fledged fewer nestlings, translating to a 5 times higher rate of population shrinkage on organic farms compared to conventional. In addition, when parental provisioning behaviour was not restricted by predation risk (i.e., at times of low corvid activity), nestling provisioning rates were higher at lower territory densities, resulting in a much greater increase in nestling mass in low density areas, suggesting that food limitation occurred at high densities. These findings in turn suggest an ecological trap, whereby preferred nesting habitat does not provide sufficient food for rearing nestlings at high population density, creating a population sink. Habitat management for farmland birds should focus not simply on creating a high nesting density, but also on ensuring heterogeneous habitats to provide food resources in close proximity to nesting birds, even if this occurs through potentially restricting overall nest density but increasing population-level breeding success.     

Pesticide Acute Toxicity Is a Better Correlate of U.S. Grassland Bird Declines than Agricultural Intensification

Common agricultural birds are in decline, both in Europe and in North America. Evidence from Europe suggests that agricultural intensification and, for some species, the indirect effects of pesticides mediated through a loss of insect food resource is in part responsible. On a state-by-state basis for the conterminous Unites States (U.S.), we looked at several agronomic variables to predict the number of grassland species increasing or declining according to breeding bird surveys conducted between 1980 and 2003. Best predictors of species declines were the lethal risk from insecticide use modeled from pesticide impact studies, followed by the loss of cropped pasture. Loss of permanent pasture or simple measures of agricultural intensification such as the proportion of land under crop or the proportion of farmland treated with herbicides did not explain bird declines as well. Because the proportion of farmland treated with insecticides, and more particularly the lethal risk to birds from the use of current insecticides feature so prominently in the best models, this suggests that, in the U.S. at least, pesticide toxicity to birds should be considered as an important factor in grassland bird declines.     

Developing indicators for European birds

The global pledge to deliver 'a significant reduction in the current rate of biodiversity loss by 2010' is echoed in a number of regional and national level targets. There is broad consensus, however, that in the absence of conservation action, biodiversity will continue to be lost at a rate unprecedented in the recent era. Remarkably, we lack a basic system to measure progress towards these targets and, in particular, we lack standard measures of biodiversity and procedures to construct and assess summary statistics. Here, we develop a simple classification of biodiversity indicators to assist their development and clarify purpose. We use European birds, as example taxa, to show how robust indicators can be constructed and how they can be interpreted. We have developed statistical methods to calculate supranational, multi-species indices using population data from national annual breeding bird surveys in Europe. Skilled volunteers using standardized field methods undertake data collection where methods and survey designs differ slightly across countries. Survey plots tend to be widely distributed at a national level, covering many bird species and habitats with reasonable representation. National species' indices are calculated using log-linear regression, which allows for plot turnover. Supranational species' indices are constructed by combining the national species' indices weighted by national population sizes of each species. Supranational, multi-species indicators are calculated by averaging the resulting indices. We show that common farmland birds in Europe have declined steeply over the last two decades, whereas woodland birds have not. Evidence elsewhere shows that the main driver of farmland bird declines is increased agricultural intensification. We argue that the farmland bird indicator is a useful surrogate for trends in other elements of biodiversity in this habitat.