Experimental safety testing confirms that the NSAID nimesulide is toxic to Gyps vultures in India.

Population declines of Gyps vultures throughout South Asia were caused by unintentional poisoning by the NSAID diclofenac, which was subsequently banned. However, other vulture-toxic NSAIDs are available, including nimesulide, which, in experiments carried out in South Africa, was shown to be toxic to Gyps vultures. We report on safety-testing of nimesulide carried out on Himalayan Griffons G. himalayensis. We gave two vultures a dose of nimesulide by oral gavage at the maximum level of exposure, with two controls dosed with benzyl alcohol. In the two tested birds, plasma nimesulide concentrations peaked after six hours, while serum uric acid concentrations increased steadily up until 24 h post-treatment, after which both birds died, displaying severe visceral gout. The control birds showed no adverse clinical or biochemical signs. We confirm that nimesulide is toxic to Gyps vultures. Veterinary use of nimesulide should be banned in all Gyps vulture range countries in the region.

Accounting for predator species identity reveals variable relationships between nest predation rate and habitat in a temperate forest songbird.

Nest predation is the primary cause of nest failure in most ground-nesting bird species. Investigations of relationships between nest predation rate and habitat usually pool different predator species. However, such relationships likely depend on the specific predator involved, partly because habitat requirements vary among predator species. Pooling may therefore impair our ability to identify conservation-relevant relationships between nest predation rate and habitat. We investigated predator-specific nest predation rates in the forest-dependent, ground-nesting wood warbler Phylloscopus sibilatrix in relation to forest area and forest edge complexity at two spatial scales and to the composition of the adjacent habitat matrix. We used camera traps at 559 nests to identify nest predators in five study regions across Europe. When analyzing predation data pooled across predator species, nest predation rate was positively related to forest area at the local scale (1000 m around nest), and higher where proportion of grassland in the adjacent habitat matrix was high but arable land low. Analyses by each predator species revealed variable relationships between nest predation rates and habitat. At the local scale, nest predation by most predators was higher where forest area was large. At the landscape scale (10,000 m around nest), nest predation by buzzards Buteo buteo was high where forest area was small. Predation by pine martens Martes martes was high where edge complexity at the landscape scale was high. Predation by badgers Meles meles was high where the matrix had much grassland but little arable land. Our results suggest that relationships between nest predation rates and habitat can depend on the predator species involved and may differ from analyses disregarding predator identity. Predator-specific nest predation rates, and their relationships to habitat at different spatial scales, should be considered when assessing the impact of habitat change on avian nesting success.

Metabolism of aceclofenac to diclofenac in the domestic water buffalo Bubalus bubalis confirms it as a threat to Critically Endangered Gyps vultures in South Asia.

Vulture declines in South Asia were caused by accidental poisoning by the veterinary non-steroidal anti-inflammatory drug (NSAID) diclofenac. Although veterinary use of diclofenac has been banned, other vulture-toxic NSAIDs are legally available, including aceclofenac, which has been shown to metabolise into diclofenac in domestic cattle. We gave nine domestic water buffalo the recommended dose of aceclofenac (2 mg kg-1 body weight), collected blood at intervals up to 48 h, and carried out a pharmacokinetic analysis of aceclofenac and its metabolite diclofenac in plasma. Aceclofenac was rapidly converted to diclofenac, and was barely detectable in plasma at any sampling time. Diclofenac was present within 20 min, and peaked 4-8 h after dosing. Aceclofenac is a prodrug of diclofenac, and behaves similarly in domestic water buffalo as it did in domestic cattle, posing the same risk to vultures. We recommend an immediate ban on the veterinary use of aceclofenac across vulture-range countries.

The non-steroidal anti-inflammatory drug nimesulide kills Gyps vultures at concentrations found in the muscle of treated cattle.

Throughout South Asia, cattle are regularly treated with non-steroidal anti-inflammatory drugs (NSAIDs) and their carcasses are left for scavengers to consume. Residues of the NSAID diclofenac in cattle carcasses caused widespread mortality and catastrophic population declines in three species of Gyps vulture during the 1990s and 2000s. Diclofenac is now banned, but other NSAIDs are used in its place. Different lines of evidence, including safety testing in Gyps vultures, have shown that some of these other NSAIDs are toxic, or probably toxic, to vultures. The NSAID nimesulide is widely available and commonly used, and has been found in dead vultures with signs of renal failure (i.e. visceral gout) and without the presence of diclofenac and/or other vulture-toxic NSAIDs. Nimesulide is therefore probably toxic to vultures. Here, we report safety testing of nimesulide in Gyps vultures. In a controlled toxicity experiment, we gave two vultures the maximum likely exposure (MLE) of nimesulide calculated from initial pharmacokinetic and residue experiments in cattle. Two other control birds were given an oral dose of water. Both vultures dosed with nimesulide died within 30 h, after showing outward signs of toxicity and increases in biochemical indicators of renal failure. Post-mortem examinations found extensive visceral gout in both vultures. Both control vultures survived without biochemical indicators of renal failure. With this evidence, we call for an immediate and comprehensive ban of nimesulide throughout South Asia to ensure the survival of the region's Critically Endangered vultures. More generally, testing the impacts of drugs on non-target species should be the responsibility of the pharmaceutical industry, before their veterinary use is licensed.

Experimental safety testing shows that the NSAID tolfenamic acid is not toxic to Gyps vultures in India at concentrations likely to be encountered in cattle carcasses.

Population declines of Gyps vultures across the Indian subcontinent were caused by unintentional poisoning by the non-steroidal anti-inflammatory drug (NSAID) diclofenac. Subsequently, a number of other NSAIDs have been identified as toxic to vultures, while one, meloxicam, is safe at concentrations likely to be encountered by vultures in the wild. Other vulture-safe drugs need to be identified to reduce the use of those toxic to vultures. We report on safety-testing experiments on the NSAID tolfenamic acid on captive vultures of three Gyps species, all of which are susceptible to diclofenac poisoning. Firstly, we estimated the maximum level of exposure (MLE) of wild vultures and gave this dose to 40 Near Threatened Himalayan Griffons G. himalayensis by oral gavage, with 15 control birds dosed with benzyl alcohol (the carrier solution for tolfenamic acid). Two birds given tolfenamic acid died with elevated uric acid levels and severe visceral gout, while the remainder showed no adverse clinical or biochemical signs. Secondly, four G. himalayensis were fed tissues from water buffaloes which had been treated with double the recommended veterinary dose of tolfenamic acid prior to death and compared to two birds fed uncontaminated tissue; none suffered any clinical effects. Finally, two captive Critically Endangered vultures, one G. bengalensis and one G. indicus, were given the MLE dose by gavage and compared to two control birds; again, none suffered any clinical effects. The death of two G. himalayensis may have been an anomaly due to i) the high dose level used and ii) the high ambient temperatures at the time of the experiment. Tolfenamic acid is likely to be safe to Gyps vultures at concentrations encountered by wild birds and could therefore be promoted as a safe alternative to toxic NSAIDs. It is manufactured in the region, and is increasingly being used to treat livestock.

Spatial behavior and habitat use in widely separated breeding and wintering distributions across three species of long-distance migrant Phylloscopus warblers.

AIM: To investigate the ecological relationship between breeding and wintering in specialist and generalist long-distance migratory species, and the links between densities and range sizes. LOCATION: Denmark, Senegal and Ghana. METHODS: We use radio tracking to study spatial behavior and habitat use in three morphologically and ecologically similar and closely related Phylloscopus species on their widely separated breeding and wintering distributions. During wintering and breeding, willow warblers P. trochilus (winter: n = 9, breeding: n = 13), chiffchaffs P. collybita (n = 11, n = 7), and wood warblers P. sibilatrix (n = 17, n = 14) were tracked. RESULTS: Willow warblers P. trochilus increased home range sizes in winter, whereas it was similar in chiffchaffs P. collybita and wood warblers P. sibilatrix, in both seasons. Home ranges overlapped more in winter than in the breeding season. In winter, home range overlap was similar among species but larger overlap during breeding was indicated for willow warblers. Tree cover was unrelated to home range size but significantly higher in breeding than in winter in all species. However, whereas willow warblers and wood warblers maintained some degree of tall tree cover inside their home ranges in winter, chiffchaffs changed from more than 80% to <1% tree cover, indicating a niche shift. MAIN CONCLUSIONS: Individuals of all three species showed changes between breeding and wintering areas in spatial behavior and habitat availability, with larger overlap in winter. The differences in patterns were potentially related to being generalist (willow warbler) or specialist (chiffchaff and wood warbler). These ecological relationships are important for the conservation of migrants and for understanding the link between breeding and wintering distributions and ecology.

Stable isotope analysis provides new information on winter habitat use of declining avian migrants that is relevant to their conservation.

Winter habitat use and the magnitude of migratory connectivity are important parameters when assessing drivers of the marked declines in avian migrants. Such information is unavailable for most species. We use a stable isotope approach to assess these factors for three declining African-Eurasian migrants whose winter ecology is poorly known: wood warbler Phylloscopus sibilatrix, house martin Delichon urbicum and common swift Apus apus. Spatially segregated breeding wood warbler populations (sampled across a 800 km transect), house martins and common swifts (sampled across a 3,500 km transect) exhibited statistically identical intra-specific carbon and nitrogen isotope ratios in winter grown feathers. Such patterns are compatible with a high degree of migratory connectivity, but could arise if species use isotopically similar resources at different locations. Wood warbler carbon isotope ratios are more depleted than typical for African-Eurasian migrants and are compatible with use of moist lowland forest. The very limited variance in these ratios indicates specialisation on isotopically restricted resources, which may drive the similarity in wood warbler populations' stable isotope ratios and increase susceptibility to environmental change within its wintering grounds. House martins were previously considered to primarily use moist montane forest during the winter, but this seems unlikely given the enriched nature of their carbon isotope ratios. House martins use a narrower isotopic range of resources than the common swift, indicative of increased specialisation or a relatively limited wintering range; both factors could increase house martins' vulnerability to environmental change. The marked variance in isotope ratios within each common swift population contributes to the lack of population specific signatures and indicates that the species is less vulnerable to environmental change in sub-Saharan Africa than our other focal species. Our findings demonstrate how stable isotope research can contribute to understanding avian migrants' winter ecology and conservation status.

Importance of climatic and environmental change in the demography of a multi-brooded passerine, the woodlark Lullula arborea.

1. We examined the influence of local weather conditions on reproductive success, timing of breeding and survival in a population of a multi-brooded ground nesting passerine (woodlark Lullula arborea) over 35 years. 2. Woodlarks laid larger clutches when rainfall was low and temperature high during the egg-laying and pre-laying period. Nest success increased with higher temperatures during the nesting period. In successful nests, the number of chicks fledged per egg laid was greater when weather was drier during the brood stage. 3. Although woodlarks bred earlier in years with warmer early spring temperatures, with the onset of breeding varying by 25 days, there was no significant advance in the onset of breeding over the 35 years of study, due to considerable inter-annual variability, and no overall trend, in weather. 4. Simulation modelling of annual reproductive output demonstrated that earlier breeding could increase productivity by 23.5% in the warmest compared to the coldest year, due to birds having more nesting attempts. Other effects of weather on productivity affected breeding output to a lesser extent. 5. Effects of weather on productivity were minor compared to an increased rate of nest predation through the period of study, which reduced productivity by 49.8% by 2004 compared to 1971. 6. Turning points analysis identified three distinct demographic periods: from 1971 to 1988 the population grew slowly, during 1988-1999 the population grew rapidly, but after 1999 the population declined. Increased population growth after 1988 was associated with higher first-year survival rates (estimated using a population model). Population decline after 1999 was caused by a combination of reduced productivity (resulting from increased nest failure rates attributed to predation) and lower first-year survival rates, that appear unrelated to winter temperature. 7. Climate change (long-term changes in weather) did not explain the marked changes observed in the population trajectory over 35 years. We suggest that understanding effects of both climate and habitat change on populations is essential in predictive population modelling.

Quantifying density dependence in a bird population using human disturbance.

Although density dependence has long been recognised as vital to population regulation, there have been relatively few studies demonstrating it spatially in wildlife populations, often due to the confounding effects of variation in habitat quality. We report on a study of woodlarks Lullula arborea, a species of European conservation concern, breeding on lowland heath in Dorset, England. We take the novel approach of utilising the birds' response to human disturbance, which resulted in much of the variation in density but had no direct impact on demographic rates. Within years, in sites with greater density there were smaller mean chick masses, lower post-fledging survival, and higher rates of nestling mortality attributed to starvation. The effects on clutch size and fledging success were confounded by the area of grassland within a site. There was no effect on brood size. Density dependence also operated within sites between years: as density increased there were reductions in mean chick mass and post-fledging survival, while nestling mortality attributed to starvation increased. Density-dependent effects on clutch size were only weakly regulatory, whereas density-dependent starvation and post-fledging mortality rates contributed strongly to differences in overall breeding output. Heavier chicks (when 7 days old) were significantly more likely to fledge and less likely to starve. Broods with heavier chicks were more likely to supply recruits to the breeding population. Nestling mass was not a factor in survival in the immediate post-fledging period, suggesting that density-dependent processes act independently on this stage. We conclude that the number of birds per hectare of suitable habitat is a valid means of expressing density, and that habitat acts as a surrogate for food abundance through which density dependence operates on the woodlark population.