Impact of the California lead ammunition ban on reducing lead exposure in golden eagles and turkey vultures

Predatory and scavenging birds may be exposed to high levels of lead when they ingest shot or bullet fragments embedded in the tissues of animals injured or killed with lead ammunition. Lead poisoning was a contributing factor in the decline of the endangered California condor population in the 1980s, and remains one of the primary factors threatening species recovery. In response to this threat, a ban on the use of lead ammunition for most hunting activities in the range of the condor in California was implemented in 2008. Monitoring of lead exposure in predatory and scavenging birds is essential for assessing the effectiveness of the lead ammunition ban in reducing lead exposure in these species. In this study, we assessed the effectiveness of the regulation in decreasing blood lead concentration in two avian sentinels, golden eagles and turkey vultures, within the condor range in California. We compared blood lead concentration in golden eagles and turkey vultures prior to the lead ammunition ban and one year following implementation of the ban. Lead exposure in both golden eagles and turkey vultures declined significantly post-ban. Our findings provide evidence that hunter compliance with lead ammunition regulations was sufficient to reduce lead exposure in predatory and scavenging birds at our study sites.     

Lead in Ammunition: A Persistent Threat to Health and Conservation

Many scavenging bird populations have experienced abrupt declines across the globe, and intensive recovery activities have been necessary to sustain several species, including the critically endangered California condor (Gymnogyps californianus). Exposure to lead from lead-based ammunition is widespread in condors and lead toxicosis presents an immediate threat to condor recovery, accounting for the highest proportion of adult mortality. Lead contamination of carcasses across the landscape remains a serious threat to the health and sustainability of scavenging birds, and here we summarize recent evidence for exposure to lead-based ammunition and health implications across many species. California condors and other scavenging species are sensitive indicators of the occurrence of lead contaminated carcasses in the environment. Transdisciplinary science-based approaches have been critical to managing lead exposure in California condors and paving the way for use of non-lead ammunition in California. Similar transdisciplinary approaches are now needed to translate the science informing on this issue and establish education and outreach efforts that focus on concerns brought forth by key stakeholders.     

Exposure of California Condors to Lead From Spent Ammunition

ABSTRACT The scientific evidence that California condors (Gymnogyps californianus) are frequently sickened and killed by lead poisoning from spent ammunition supports the conclusion that current levels of lead exposure are too high to allow reintroduced condors to develop self-sustaining populations in the wild in Arizona and, by inference, in California. The evidence for lead poisoning and its source comes from the following sorts of data: 1) 18 clinical necropsies revealing high levels of lead in body tissues and (or) presence of lead shotgun pellets and bullet fragments in digestive tracts; 2) moribund condors showing crop paralysis and impending starvation with toxic levels of lead in their blood; 3) widespread lead exposure among free-flying condors, many with clinically exposed or acute levels; 4) temporal and spatial correlations between big game hunting seasons and elevated lead levels in condors; and 5) lead isotope ratios from exposed condors showing close similarity to isotope ratios of ammunition lead but isotope ratios in less exposed condors being similar to environmental background sources, which are different from ammunition lead. Simple population models reveal harmful demographic impacts of unnatural mortality from lead on population trajectories of reintroduced condors. Recent innovations in the manufacture of nonlead shotgun pellets and bullets with superior ballistics now provide for a simple solution to the problem of lead ingestion by condors, many other species of wildlife, and human beings: substitute nontoxic forms of ammunition for traditional lead-based ammunition. The substitution of nontoxic ammunition would be highly efficacious for hunting, economically feasible, and the right thing to do.     

Determining prevalence of bluetongue and epizootic hemorrhagic disease viruses in mule deer in Arizona (USA) using whole blood dried on paper strips compared to serum analyses

We investigated the feasibility of using whole blood dried on paper strips as a means to collect antibody prevalence data for the epizootic hemorrhagic disease viruses (EHDV) and bluetongue viruses (BTV) from hunter-harvested male mule deer (Odocoileus hemionus) in October 2002 from Arizona, USA. We compared antibody prevalence estimates in mule deer from paired paper strip and serum samples. Prevalence data obtained from elution of dried blood on paper strips proved to be consistent with results from serum in 94% of the samples tested. The paper strip method allows easy collection of blood from dead animals, with a smaller amount of blood being needed for analyses. Also, samples do not need to be refrigerated before analyses. We also used serum samples to determine hemorrhagic disease (HD) serotype exposure status of mule deer harvested from 4 distinct areas in Arizona. Antibodies to BTV and EHDV were identified in 3 of the 4 areas, with positive results to EHDV-1, EHDV-2, BTV-10, and BTV-11 being most common. Many animals did not have antibodies against the BTV serotypes. Exposure varied geographically and potentially with elevation. Hemorrhagic disease viruses commonly infect Arizona mule deer, except on the Kaibab Plateau in northern Arizona.     

Lead exposure in free-flying turkey vultures is associated with big game hunting in California

Predatory and scavenging birds are at risk of lead exposure when they feed on animals injured or killed by lead ammunition. While lead ammunition has been banned from waterfowl hunting in North America for almost two decades, lead ammunition is still widely used for hunting big game and small game animals. In this study, we evaluated the association between big game hunting and blood lead concentration in an avian scavenger species that feeds regularly on large mammals in California. We compared blood lead concentration in turkey vultures within and outside of the deer hunting season, and in areas with varying wild pig hunting intensity. Lead exposure in turkey vultures was significantly higher during the deer hunting season compared to the off-season, and blood lead concentration was positively correlated with increasing wild pig hunting intensity. Our results link lead exposure in turkey vultures to deer and wild pig hunting activity at these study sites, and we provide evidence that spent lead ammunition in carrion poses a significant risk of lead exposure to scavengers.     

Reducing Lead on the Landscape: Anticipating Hunter Behavior in Absence of a Free Nonlead Ammunition Program

Lead is a neurotoxin that has been documented to affect many forms of wildlife, and has been identified as a limiting factor in a population of California Condors in Northern Arizona. The Arizona Game and Fish Department provides vouchers for free nonlead ammunition to hunters selected to hunt within the distribution of California Condors, with the intention of having fewer lead-laden offal piles available to California Condors. Although wildlife agencies may reasonably assume voucher programs motivate hunters into choosing nonlead ammunition, the lead reduction efforts attributable to the voucher program has not been empirically quantified. Our intention was to compare a control group of hunters to a treatment group of hunters within California Condor occupied areas. Both groups received educational materials regarding the deleterious effects of lead, but the treatment group also received a voucher for a free initial box of ammunition. About half of the control group used nonlead ammunition, compared to about three-fourths of the treatment group. Prominent barriers to adoption of nonlead ammunition included a general difficulty of obtaining it, obtaining it in the desired caliber, and its costliness. Frequently mentioned motivations for using nonlead was the exhortation to use it by the Department, and the desire to aid California Condor recovery by hunters. The disparate compliance rates found herein confirm and quantify the success of nonlead ammunition voucher programs, but underscore the importance of working to increase the supply of nonlead ammunition with the end of facilitating its procurement and reducing its cost.     

Evidence of High Tolerance to Ecologically Relevant Lead Shot Pellet Exposures by an Upland Bird

Upland birds that display grit ingestion behavior are potentially at risk of detrimental effects and death from lead poisoning at trap and skeet ranges and other areas where vast quantities of spent lead shot pellets abound. Because commonly cited force-feeding pellet exposure studies deviate from true field conditions, their results may not reflect true risks faced by upland birds. In particular, studies that use new shot pellets and administer more pellets than would be reasonably ingested, critically interfere with the understanding of actualized pellet exposures. In this study, northern bobwhites (Colinus virginianus), a frequent test species in shot pellet research, were dosed in an ecologically-relevant manner (i.e., with spent shot and with no more than three pellets). Notably, the 56-day post-dosing observation period, during which a battery of physiological measures were recorded, exceeded that of related studies. Despite a sustained suppression of a lead poisoning indicator, the data suggest upland birds can withstand spent shot pellet exposures. Data detected a survivorship ≥95%, absence of illness, demonstrated tolerance for extremely high blood lead concentrations, and unaffected blood parameters. In conjunction with ecological considerations (e.g., spatial scale and animal behavior), concern about bird population losses from the incidental ingestion of spent shot pellets is potentially overstated.     

Experimental lead poisoning in turkey vultures (Cathartes aura)

Lead-induced mortality appears to have been a major factor in the decline of the California condor (Gymnogyps californianus). We orally dosed turkey vultures (Cathartes aura) with BB-sized lead shot from January 1988 through July 1988 to determine physiologic response (delta-aminolevulinic acid dehydratase inhibition, erythrocyte protoporphyrin levels, anemia), diagnostic tissue lead concentrations (blood, liver, and kidney), and comparative sensitivity of this species. Two turkey vultures died and two became so intoxicated they were euthanized. Overall, responses of measured parameters were comparable to other species exposed to lead although there was considerable individual variation. Survival time (143-211 days), even with the large numbers of shot and constant redosing, was much longer than reported for other species of birds, suggesting considerable tolerance by turkey vultures to the deleterious effects of lead ingestion. Based on these observations, turkey vultures appear to be poor models for assessing the risk of lead poisoning to California condors or predicting their physiologic response.     

Sources of Mortality in Bald Eagles in Michigan, 1986–2017

As bald eagle populations recover, defining major sources of mortality provides managers important information to develop management plans and mitigation efforts. We obtained data from necropsies on 1,490 dead bald eagles (Haliaeetus leucocephalus) collected in Michigan, USA, conducted from 1986 to 2017 to determine causes of death (COD). Trauma and poisoning were the most common primary COD categories, followed by disease. Within trauma and poisoning, vehicular trauma (n = 532) and lead poisoning (n = 176) were the leading COD subcategories, respectively. Females comprised a greater number of carcasses for most COD diagnoses. The proportion of trauma and poisoning CODs significantly increased in the last few years of the study in comparison to a select few years at the beginning. Trauma CODs were greater in autumn months during whitetail deer (Odocoileus virginianus) breeding and hunting seasons and in February, when aquatic foraging is unavailable and eagles are likely forced to scavenge along roadsides. Poisoning CODs were greatest in late winter and early spring months, when deer carcasses containing lead ammunition, which are preserved by the cold weather, also become a supplemental food source. The major infectious disease CODs, West Nile virus and botulism (Clostridium botulinum type E), were more prevalent during summer months. We recommend moving road-killed carcasses, especially white-tailed deer, from the main thoroughfare to the back of the right-of-way, and the transition from lead ammunition and fishing tackle to non-toxic alternatives to decrease these main anthropogenic sources of mortality for bald eagles, and other scavenger species. © 2020 The Wildlife Society.     

Lead poisoning in swans and sources of contamination in Ireland

A study of blood levels in mute swans Cygnus olor in Ireland has revealed that ingested lead pellets are responsible for acute lead poisoning. Forty-two percent of blood samples from 890 live birds at one site showed elevated lead levels. X-ray examination of live birds revealed the source of contamination to be ingested lead pellets. Urban birds were shown to have higher ( P 0001) lead levels than rural birds, the blood lead levels of which were presumed to reflect natural background levels. Urban grass was shown to have elevated lead but this did not cause lead poisoning in Canada geese Branta canadensis . Post-mortem examination has shown that 68% ( n = 101) of all mute swans examined from a number of sites died from lead poisoning. Two sources of poisoning were identified; spent gunshot from a claypigeon shooting site at Lough Neagh, Northern Ireland, and lost or discarded anglers' weights at Cork Lough and at a fishing pond in Belfast, N.I. The first known case of lead poisoning in whooper swans Cygnus cygnus in Ireland is recorded which resulted from the ingestion of gunshot used almost two decades earlier. Aspects of the pathology of lead poisoned swans is discussed.     

Ammunition Type and Shot Placement Determine Lead Fragmentation in Deer

Lead remnants from spent ammunition introduce health risks to humans and wildlife that consume game harvested with firearms. Most current research has focused on effects of high-velocity rifle bullets, whereas low-velocity lead ammunition has received little attention. We examined whether fragmentation characteristics differed between 3 common low-velocity ammunition types when shot into the thoracic cavity or shoulder of white-tailed deer (Odocoileus virginianus) culled in Indiana, USA, from May–September of 2009. We shot and subsequently radiographed 43 deer to determine the number of fragments, fragment size, and distance traveled by individual fragments. We also radiographed deer post-evisceration to determine the proportion of fragments available to humans and wildlife scavengers for consumption in muscle and visceral tissue, respectively. All radiographed deer had evidence of fragmentation, with a geometric mean of 13.1 (95% CI = 10.3, 16.8) fragments/deer. Most fragments (89%) were <5 mm from wound channels, and no fragment traveled beyond 205 mm from a wound channel. Fragments were often retained within the muscle tissue of deer with a geometric mean rate of 0.55 (95% CI = 0.48, 0.65). Muzzleloader bullet fragments were larger than those generated by rifled and sabot slugs, and sabot slug fragments had the shortest dispersal from wound channels. Shoulder-shot placement and bone contact for all ammunition resulted in a greater number of fragments (P < 0.01). Shoulder-shots also generated more small fragments and higher fragment retention in muscle tissue. The overall mean number of lead fragments detected across our ammunition treatments was less than previous studies. Our results indicate ammunition type and shot placement may be considerations for hunters wishing to limit their potential exposure to lead from harvested big game. Additionally, when compared to high-velocity rifle bullets, significantly fewer lead fragments are made available to humans and wildlife that consume game harvested with low-velocity ammunition types, such as those tested here. Complete elimination of lead ingestion by humans and wildlife, however, is only possible by using non-toxic ammunition alternatives. © 2020 The Wildlife Society.     

Cytogenetic analysis of California condor (Gymnogyps californianus) chromosomes: comparison with chicken (Gallus gallus) macrochromosomes

The California condor is the largest flying bird in North America and belongs to a group of New World vultures. Recovering from a near fatal population decline, and currently with only 197 extant individuals, the species remains listed as endangered. Very little genetic information exists for this species, although sexing methods employing chromosome analysis or W-chromosome specific amplification is routinely applied for the management of this monomorphic species. Keeping in mind that genetic conditions like chondrodystrophy have been identified, preliminary steps were undertaken in this study to understand the genome organization of the condor. This included an extensive cytogenetic analysis that provided (i) a chromosome number of 80 (with a likelihood of an extra pair of microchromosomes), and (ii) information on the centromeres, telomeres and nucleolus organizer regions. Further, a comparison between condor and chicken macrochromosomes was obtained by using individual chicken chromosome specific paints 1-9 and Z and W on condor metaphase spreads. Except for chromosomes 4 and Z, each of the chicken (GGA) macrochromosomes painted a single condor (GCA) macrochromosome. GGA4 paint detected complete homology with two condor chromosomes, viz., GCA4 and GCA9 providing additional proof that the latter are ancestral chromosomes in the birds. The chicken Z chromosome showed correspondence with both Z and W in the condor. The homology suggests that the condor sex chromosomes have not completely differentiated during evolution, which is unlike the majority of the non-ratites studied up till now. Overall, the study provides detailed cytogenetic and basic comparative information on condor chromosomes. These findings significantly advance the effort to study the chondrodystrophy that is responsible for over ten percent mortality in the condor.     

Energy beyond food: foraging theory informs time spent in thermals by a large soaring bird

Current understanding of how animals search for and exploit food resources is based on microeconomic models. Although widely used to examine feeding, such constructs should inform other energy-harvesting situations where theoretical assumptions are met. In fact, some animals extract non-food forms of energy from the environment, such as birds that soar in updraughts. This study examined whether the gains in potential energy (altitude) followed efficiency-maximising predictions in the world's heaviest soaring bird, the Andean condor (Vultur gryphus). Animal-attached technology was used to record condor flight paths in three-dimensions. Tracks showed that time spent in patchy thermals was broadly consistent with a strategy to maximise the rate of potential energy gain. However, the rate of climb just prior to leaving a thermal increased with thermal strength and exit altitude. This suggests higher rates of energetic gain may not be advantageous where the resulting gain in altitude would lead to a reduction in the ability to search the ground for food. Consequently, soaring behaviour appeared to be modulated by the need to reconcile differing potential energy and food energy distributions. We suggest that foraging constructs may provide insight into the exploitation of non-food energy forms, and that non-food energy distributions may be more important in informing patterns of movement and residency over a range of scales than previously considered.     

Patterns of mortality in free-ranging California Condors (Gymnogyps californianus)

We document causes of death in free-ranging California Condors (Gymnogyps californianus) from the inception of the reintroduction program in 1992 through December 2009 to identify current and historic mortality factors that might interfere with establishment of self-sustaining populations in the wild. A total of 135 deaths occurred from October 1992 (the first post-release death) through December 2009, from a maximum population-at-risk of 352 birds, for a cumulative crude mortality rate of 38%. A definitive cause of death was determined for 76 of the 98 submitted cases, 70% (53/76) of which were attributed to anthropogenic causes. Trash ingestion was the most important mortality factor in nestlings (proportional mortality rate [PMR] 73%; 8/11), while lead toxicosis was the most important factor in juveniles (PMR 26%; 13/50) and adults (PMR 67%; 10/15). These results demonstrate that the leading causes of death at all California Condor release sites are anthropogenic. The mortality factors thought to be important in the decline of the historic California Condor population, particularly lead poisoning, remain the most important documented mortality factors today. Without effective mitigation, these factors can be expected to have the same effects on the sustainability of the wild populations as they have in the past.     

Comparison of lead contamination among related scavenging birds sharing food sources: a call for caution when using surrogate species

Lead contamination is a global problem affecting a large number of bird species around the world. Among the different avian guilds, vultures and facultative scavengers are particularly threatened by this toxic metal. However, little information is available about differences in exposure to this metal for sympatric vulture species that share food resources. We compared blood lead concentrations of two closely related sympatric obligate scavenger species, the abundant Black Vulture Coragyps atratus and the threatened Andean Condor Vultur gryphus in north-western Patagonia, Argentina. We sampled 28 Andean Condors and 29 Black Vultures trapped foraging in the same area in the steppe. We also sampled 16 Black Vultures foraging in a rubbish dump to determine whether there were differences in lead contamination among foraging sites. Andean Condors had significantly higher mean blood lead concentrations than Black Vultures. There was no difference in lead concentrations between Black Vultures trapped in the steppe and in the rubbish dump. The prevalence and probability of lead concentrations above the threshold level (20 µg/dL) was higher for Andean Condors than for Black Vultures, potentially producing different effects on their health. This disparity in lead contamination may be due to differences in their foraging habits or in their susceptibility to this toxic metal. Overall, our results suggest caution in using an abundant surrogate species to infer lead contamination in a closely related but harder to sample species.     

Lead poisoning in wild populations of Marsh Harriers Circus aeruginosus in the Camargue and Charente-Maritime, France

Lead poisoning of waterfowl, through the ingestion of spent gunshot, has been recognized as a mortality factor for over a century. However, in Europe relatively little attention has been paid to raptors that may ingest shot embedded in the flesh of prey. The present study examines the incidence of lead poisoning in wild Marsh Harriers Circus aeruginosus trapped at two sites in France, the Camargue and Charente-Maritime.
Eight captive Marsh Harriers (controls) had blood lead (PbB) concentrations of 5.3–10.8 μg per dl. Of 94 wild birds trapped during the winters of 1990/1991 and 1991/1992 either in baited clap traps or in mist nets at night roosts, 29 (31%) had elevated (>30 μg/dl) PbB concentrations and 13 (14%) had concentrations indicative of clinical poisoning (> 60 μg/dl). Similar percentages of birds caught using the two trapping methods had elevated (>30 μg/dl) PbB concentrations. However, clap netting appeared to select for more heavily contaminated birds and for juvenile birds, although the two are not necessarily correlated. In the Camargue, significantly more females than males trapped in clap nets had elevated PbB concentrations.
The incidence of shot in regurgitated Marsh Harrier pellets increased significantly between October and December, indicating increased exposure to lead as the hunting season progressed. There appeared to be a parallel increase in PbB concentrations throughout the hunting season, although this could not be clearly demonstrated as sex ratios were different during different sampling periods.
Elevated PbB concentrations of harriers in the Camargue are likely to result primarily from the ingestion of shot in the flesh of crippled or unretrieved waterfowl and in Charente-Maritime, from eating crippled or dead mammals.
Other raptors at risk from lead poisoning and solutions to this problem are discussed.     

Was Aldo Leopold Right about the Kaibab Deer Herd?

In ecology textbooks prior to the 1970s, Aldo Leopold’s classic story of predator control, overpopulation of deer, and habitat degradation on the Kaibab Plateau during the 1920s epitomized predator regulation of herbivore populations. However, the story disappeared from texts in the late 20th century after several papers noted uncertainties in estimations of the deer population and provided alternative explanations. We re-examined the case study by determining the age structure of aspen (Populus tremuloides Michx.) on the plateau. Aspen comprises the majority of deer browse in the summer, and the absence of a normal cohort of aspen from the 1920s would indicate deer overpopulation. The number of aspen (at 1.4 m) dating to the 1920s was an order of magnitude lower than the null expectation. Other periods of unusual numbers of aspen included high numbers of aspen dating to the 1880s and 1890s (when regular surface fires ceased), few aspen dating from 1953 to 1962 (after a second irruption of the deer population), and very high numbers from 1968 to 1992 (coincident with widespread logging). These convergent lines of evidence support the idea of extreme deer herbivory in the 1920s, consistent with food limitation of deer at high populations (bottom–up control) and predation limitation at low deer populations (top–down control). Some uncertainty remains within the overall story, and this level of ambiguity is common in case studies that involve population ecology, land management, and people at the scale of 1,000 km² and 100 years. A complete version of the Kaibab deer story and its history would be a valuable, realistic case study for ecology texts.     

Adverse health effects in Canada geese (Branta canadensis) associated with waste from zinc and lead mines in the Tri-State Mining District (Kansas, Oklahoma, and Missouri, USA)

Lead and zinc poisoning have been recorded in a variety of bird species, including migrating waterfowl such as Canada Geese (Branta canadensis), at sites contaminated with mine waste from lead and zinc mines in the Tri-State Mining District, Kansas, Oklahoma, and Missouri, USA. The adverse health impacts from mine waste on these birds may, however, be more extensive than is apparent from incidental reports of clinical disease. To characterize health impacts from mine waste on Canada Geese that do not have observable signs of poisoning, four to eight apparently healthy birds per site were collected from four contaminated sites and an uncontaminated reference site, and examined for physical and physiologic evidence of metals poisoning. Tissue concentrations of silver, aluminum, arsenic, barium, cadmium, cobalt, chromium, copper, iron, magnesium, manganese, molybdenum, nickel, lead, selenium, thallium, vanadium, and zinc were determined by inductively coupled plasma mass spectroscopy. Adverse health effects due to lead were characterized by assessing blood δ-aminolevulinic acid dehydratase (ALAD) enzyme activity. Adverse effects associated with zinc poisoning were determined from histologic examination of pancreas tissues. Elevated tissue lead concentrations and inhibited blood ALAD enzyme activities were consistently found in birds at all contaminated sites. Histopathologic signs of zinc poisoning, including fibrosis and vacuolization, were associated with elevated pancreatic zinc concentrations at one of the study sites. Adverse health effects associated with other analyzed elements, or tissue concentrations indicating potentially toxic exposure levels to these elements, were not observed.     

Seroepidemiologic Survey of Potential Pathogens in Obligate and Facultative Scavenging Avian Species in California

Throughout the world, populations of scavenger birds are declining rapidly with some populations already on the brink of extinction. Much of the current research into the factors contributing to these declines has focused on exposure to drug residues, lead, and other toxins. Despite increased monitoring of these declining populations, little is known about infectious diseases affecting scavenger bird species. To assess potential infectious disease risks to both obligate and facultative scavenger bird species, we performed a serosurvey for eleven potential pathogens in three species of scavenging birds in California: the California condor (Gymnogyps californianus), turkey vulture (Cathartes aura) and golden eagle (Aquila chrysaetos). California condors were seropositive for avian adenovirus, infectious bronchitis virus, Mycoplasma gallisepticum, avian paramyxovirus-2, West Nile virus (WNV) and Toxoplasma gondii. Golden eagles were seropositive for avian adenovirus, Chlamydophila psittaci and Toxoplasma gondii, and turkey vultures were seropositive for avian adenovirus, Chlamydophila psittaci, avian paramyxovirus-1, Toxoplasma gondii and WNV. Risk factor analyses indicated that rearing site and original release location were significantly associated with a positive serologic titer to WNV among free-flying condors. This study provides preliminary baseline data on infectious disease exposure in these populations for aiding in early disease detection and provides potentially critical information for conservation of the endangered California condor as it continues to expand its range and encounter new infectious disease threats.     

Use of grit supplements by waterbirds: an experimental assessment of strategies to reduce lead poisoning

The ingestion of spent Pb shot due to confusion with grit or inadvertently with food particles causes Pb poisoning in a large number of waterbirds, this being one of the main causes of mortality for some species. Lead ammunition for hunting is being progressively banned in more countries, while grit supplementation has been proposed as a management measure to reduce the ingestion of deposited Pb shot. Studies of grit selection with waterfowl in semi-captivity and in the wild were undertaken to evaluate preferences in the colour and geochemical composition of grit, whether it was available dry or in water, its position within the wetland and the relationship between grit ingestion and feeding behaviour. Grit ingestion was higher when food was included in the treatments. In the absence of food, red grit was taken in higher amounts than grey in semi-captivity but not in the wild. Siliceous grit was taken in a higher amount than calcareous when offered dry, but not in water. No differences in the amount of ingested grit were found among different positions within the wetland. The number of feeding attempts in plots supplemented with grit was higher than in those without grit, although the highest numbers of feeding birds were found in plots supplemented with food. Grit ingestion in waterfowl is intimately associated with feeding behaviour. To optimize the effectiveness of grit supplementation to reduce the risk of Pb poisoning in waterfowl, calcareous and siliceous grit may be combined and applied in feeding sites or mixed with bait to attract birds.