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1.
In several seabirds, nutritional state of a nest-bound chick is negatively correlated with the activity of its hypothalamus–pituitary–adrenal (HPA) axis. Increased corticosterone (cort) secretion has been shown to facilitate changes in behavior that allow hungry chicks to obtain more food from parents. However, if parents are not willing/able to buffer their young from temporary food shortages, increased cort secretion could be detrimental to undernourished chicks. In a system where parents are insensitive to chick demands, low benefits and high costs of activation of the HPA-axis in hungry chicks should lead to a disassociation of the nutritional state of the young and the activity of its HPA-axis. We tested this novel hypothesis for the tufted puffin (Fratercula cirrhata), a seabird with intermittent provisioning of a nest-bound semi-precocial chick. We examined the HPA-axis activity of captive chicks exposed to the following: (1) a short-term (24 h) food deprivation; and (2) an array of prolonged (3 weeks) restrictions in feeding regimens. We found that in response to a short-term food deprivation chicks decreased baseline levels of cort and thyroid hormones. In response to prolonged restrictions, food-limited chicks exhibited signs of nutritional deficit: they had lower body mass, endogenous lipid reserves, and thyroid hormone titers compared to chicks fed ad libitum. However, baseline and maximum acute stress-induced levels of cort were also lower in food-restricted chicks compared to those of chicks fed ad libitum. These results support a major prediction of the study hypothesis that puffin chicks suppress HPA-axis activity in response to short- and long-term nutritional deficits. This physiological adaptation may allow a chick to extend its development in the nest, while eluding detrimental effects of chronic cort elevation.  相似文献   

2.
Relating the effects of foraging niche variation to reproductive dynamics is critical to understand species response to environmental change. We examined foraging niche variations of the slender‐billed gull (Chroicocephalus genei), a nomadic colonial waterbird species during its range expansion along the French Mediterranean coast over a 16‐year period (1998–2013). We investigated whether range expansion was associated with a change in chick diet, breeding success, and chicks body condition. We also examined whether breeding success and chicks body condition were explained by diet and colonial characteristics (number of pairs, laying phenology, habitat, and locality). Diet was characterized using dual‐stable isotopic proxies (δ13C and δ15N) of feather keratin from 331 individuals subsampled from a total of 4,154 chicks ringed and measured at 18 different colonies. δ13C decreased and δ15N increased significantly during range expansion suggesting that chicks were fed from preys of increasing trophic level found in the less salty habitat colonized by the end of the study period. Niche shift occurred without significant change of niche width which did not vary among periods, habitats, or localities either. Breeding success and chick body condition showed no consistent trends over years. Breeding success tended to increase with decreasing δ13C at the colony level while there was no relationship between stable isotope signatures and chick body condition. Overall, our results suggest that even if range expansion is associated with foraging niche shift toward the colonization of less salty and more brackish habitats, the shift had marginal effect on the breeding parameters of the Slender‐billed gull. Niche width appears as an asset of this species, which likely explains its ability to rapidly colonize new locations.  相似文献   

3.
We studied the regulation of provisioning in Cory's shearwaterat Selvagem Grande during the chick rearing period. Provisioningwas examined in terms of feeding frequency and amount of fooddelivered to chicks. Two groups of chicks were subjected toshort-term contrasting manipulations of their nutritional status:one group of chicks was given a food supplement of about 30g, and another group was deprived of up to 30 g of food. Adultstending deprived chicks increased the frequency of feedingvisits (but not the size of feeds), which resulted in an increasein the net rate of food delivery. At the end of this study,deprived chicks were growing at the same rate as fed chicks. Parents attending fed chick did not change their provisioningrates in response to the treatment. Our results indicate thatCory's shearwaters are able to adjust their provisioning ratein response to short-term variation in the nutritional statusof their chicks. We also examined the change in the beggingrate of fed and deprived chicks in response to the treatment.There was no relationship between the begging rate and thecondition of chicks, which is taken to be a measure of thechick's physiological condition, related to its ability towithstand imposed periods of fasting. However, fed chicks decreasedtheir begging rate after the increase in their condition dueto supplementary food. Conversely, deprived chicks, which wereonly able to sustain their condition before the onset of thetreatment, maintained high levels of begging. To some extent,these results suggest that parental provisioning can be influencedby the begging behavior of chicks.  相似文献   

4.
A central point in life history theory is that parental investment in current reproduction should be balanced by the costs in terms of residual reproductive value. Long-lived seabirds are considered fixed investors, that is, parents fix a specific level of investment in their current reproduction independent to the breeding requirements. We tested this hypothesis analysing the consequences of an experimental increase in flying costs on the foraging ecology, body condition and chick condition in Cory’s shearwaters Calonectris diomedea. We treated 28 pairs by reducing the wing surface in one partner and compared them with 14 control pairs. We monitored mass changes and incubation shifts and tracked 19 foraging trips per group using geolocators. Furthermore, we took blood samples at laying, hatching and chick-rearing to analyse the nutritional condition, haematology, muscle damage and stable isotopes. Eighty-day-old chicks were measured, blood sampled and challenged with PHA immune assay. In addition, we analysed the effects of handicap on the adults at the subsequent breeding season. During incubation, handicapped birds showed a greater foraging effort than control birds, as indicated by greater foraging distances and longer periods of foraging, covering larger areas. Eighty-day-old chicks reared by treated pairs were smaller and lighter and showed a lower immunity than those reared by control pairs. However, oxygen demands, nutritional condition and stable isotopes did not differ between control and handicapped birds. Although handicapped birds had to increase their foraging effort, they maintained physical condition by reducing parental investment and transferred the experimentally increased costs to their partners and the chick. This result supports the fixed investment hypothesis and is consistent with life history theory.  相似文献   

5.
In 2002 and 2003, blood samples from white stork (Ciconia ciconia) chicks were examined for the presence of antibodies against Listeria monocytogenes. Listeria antibodies were detected in 121 (59%) of 205 chick samples. The probability of Listeria antibodies being present increased with chick age; chicks detected with Listeria antibodies were in better condition than those without the bacterium.  相似文献   

6.
Parents of albatross and shearwater species employ a dual foraging strategy, feeding their chicks quickly in repeated short trips and then restoring their own fuel reserves during longer trips. A decline in parental body condition is believed to trigger longer trips, but chick body condition and age may also play a role. To investigate these factors in the little-studied streaked shearwater Calonectris leucomelas, we monitored the nest attendance of 17 pairs on Mikura Island in 2005 using an automated identification system. We also monitored body mass changes and meal masses of 5 of the 17 pairs using an automated weighing system. Although the birds did not show a clear dual foraging pattern, trip duration varied widely from 1 to 15 days. On average, the birds fed chicks 67.6 g during nighttime meals at 2.74-day intervals. Since meal mass did not depend on trip duration, feeding efficiency (meal mass delivered per unit trip duration) decreased as trip duration increased. Parents accumulated more energy reserves when they took longer trips. Parents appeared likely to initiate longer trips when their body condition declined or chick body condition recovered.  相似文献   

7.
We investigated the seasonal pattern of changes in body condition of breeding black‐legged kittiwakes Rissa tridactyla in Svalbard (79°N) to evaluate whether changes in body condition were a consequence of the energetic demands of breeding (the reproductive stress hypothesis) or of voluntary anorexia to attain lower flight costs during chick rearing (the programmed anorexia hypothesis). Adult body condition was recorded from early egg laying to fledging and was examined in relation to date (relative to hatching), sex, parental time‐budget, brood size and reproductive success. To distinguish between the two hypotheses we evaluate whether the reduction in body condition occurred during or ahead of the energetically most demanding part of the chick‐rearing period. We combine our results on changes in body condition and time‐budget with published information on field metabolic rate (FMR) and chick energy requirements from studies in the same colony.
Our calculations of adult energy requirements and energy intakes indicate that the first part of the chick‐rearing period was energetically the most demanding period, because adult energy requirement per hour spent off the nest was highest in this period, and adults were time constrained because of the need for 24‐h brooding of the chicks.
During the incubation period female body condition increased slightly, but significantly, while male body condition was stable. During the first part of the chick‐rearing period, female and male body condition decreased by 14.8% and 8.4%, respectively. During the second part of the chick‐rearing period, both male and female body condition remained stable. The reductions in body condition occurred during the phase which was suggested to be the energetically most demanding part of the chick‐rearing period, thus supporting the reproductive stress hypothesis. Parental body condition during the incubation period was positively related to the probability of successfully fledging young, providing additional support for the reproductive stress hypothesis.  相似文献   

8.
We studied the behavioral development of seven lesser adjutant stork (Leptoptilos javanicus) chicks from hatching to fledging over three breeding seasons at the Bronx Zoo. We developed an ethogram and compared the rate at which behaviors appeared in relation to brood size, sex, and the conditions in which the chicks were raised by their parents. Although sample sizes were small, there seem to be sex‐related differences in the rate at which behaviors develop, with females developing more rapidly than males. Larger clutch size may be associated with slower growth rate because the single male developed faster than the two males in 2004. The slowest growth rate, observed in a single male chick in 1999, was most likely owing to nutritional deficiencies and other health complications. More research is needed, but these results can be used to help evaluate the age and health of lesser adjutant stork chicks in captivity and in the wild. Zoo Biol 26:533–538, 2007. © 2007 Wiley‐Liss, Inc.  相似文献   

9.
The heat increment of feeding (HIF), a transient postprandial increase in metabolic rate, is the energy cost of processing a meal. We measured HIF in house wren chicks (Troglodytes aedon) ranging in mass from 1.6 to 10.3 g. This mass range (age 2–10 days) spanned a transition from blind, naked, ectothermic chicks through alert, endothermic birds with nearly complete feathering. We fed chicks crickets (2.7–10% of chick body mass) and determined HIF from continuous measurements of oxygen consumption rate (O2) before and after meals. At warm ambient temperatures (T a) of 33–36 °C, the magnitude of HIF (in ml O2 or joules) was linearly related to meal mass and was not affected by chick mass. HIF accounted for 6.3% of ingested energy, which is within the range of results for other carnivorous vertebrates. The duration of HIF was inversely related to chick mass; 10-g chicks processed a standard meal approximately twice as fast as 2-g chicks. HIF duration increased with increasing meal mass. The peak O2 during HIF, expressed as the factorial increase above resting metabolism, was independent of body mass and meal mass. In large, endothermic chicks ( > 8 g), HIF substituted for thermoregulatory heat production at low T a. Accepted: 11 December 1996  相似文献   

10.
Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.  相似文献   

11.
Behavioral and/or developmental plasticity is crucial for resisting the impacts of environmental stressors. We investigated the plasticity of adult foraging behavior and chick development in an offshore foraging seabird, the black noddy (Anous minutus), during two breeding seasons. The first season had anomalously high sea-surface temperatures and ‘low’ prey availability, while the second was a season of below average sea-surface temperatures and ‘normal’ food availability. During the second season, supplementary feeding of chicks was used to manipulate offspring nutritional status in order to mimic conditions of high prey availability. When sea-surface temperatures were hotter than average, provisioning rates were significantly and negatively impacted at the day-to-day scale. Adults fed chicks during this low-food season smaller meals but at the same rate as chicks in the unfed treatment the following season. Supplementary feeding of chicks during the second season also resulted in delivery of smaller meals by adults, but did not influence feeding rate. Chick begging and parental responses to cessation of food supplementation suggested smaller meals fed to artificially supplemented chicks resulted from a decrease in chick demands associated with satiation, rather than adult behavioral responses to chick condition. During periods of low prey abundance, chicks maintained structural growth while sacrificing body condition and were unable to take advantage of periods of high prey abundance by increasing growth rates. These results suggest that this species expresses limited plasticity in provisioning behavior and offspring development. Consequently, responses to future changes in sea-surface temperature and other environmental variation may be limited.  相似文献   

12.
Abstract: Past studies suggest that the productivity of common loons (Gavia immer) is lower on acidic lakes in northern Wisconsin, USA, than on neutral lakes. Two hypotheses have been proposed to explain low chick survival: (1) reduced food consumption related to changes in prey communities on lower pH lakes and (2) high mercury (Hg) exposure on lower pH lakes. To address these hypotheses, we quantified prey and Hg consumption by loon chicks on 51 lakes and survival on 55 lakes ranging in pH from 4.9 to 9.5 in northern Wisconsin in 1995 and 1996. The time adults spent providing prey to chicks was unrelated to lake pH but increased with number of chicks and chick age. The number of prey caught per provisioning time declined as lake pH declined because adults made fewer dives, not because success of prey capture declined. Chicks consumed more insect larvae on acidic lakes and more crayfish (Family Astacidae) on neutral lakes. Biomass consumed ranged from an average 1.99 ± 1.05 (SE) g/hr/chick during the first week of a chick's life to a peak of 7.93 ± 1.93 g/hr/chick during the eighth week. Biomass intake per chick body weight (g/Wg/hr) declined with lake acidity but was not related to chick survival (P = 0.25). Although the Hg concentration in the 3 major prey species was positively related to lake acidity and blood Hg level of chicks at a lake, total Hg consumption (μg/Wg/hr) was highest on moderately acidic lakes rather than on the most acidic lakes. We suggest that loon chick survival in northern Wisconsin lakes is more likely related to prey availability than to Hg exposure. When we removed from our analysis 1 lake where 2 11-day-old chicks were killed by predators, chick survival was negatively related to lake acidity but not to biomass or Hg consumption. We discuss mechanisms of Hg excretion that may allow young chicks to survive on acidic lakes in northern Wisconsin despite high Hg intake.  相似文献   

13.
Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.  相似文献   

14.
ABSTRACT For seabirds raising young under conditions of limited food availability, reducing chick provisioning and chick growth rates are the primary means available to avoid abandonment of a breeding effort. For most seabirds, however, baseline data characterizing chick growth and development under known feeding conditions are unavailable, so it is difficult to evaluate chick nutritional status as it relates to foraging conditions near breeding colonies. To address this need, we examined the growth and development of young Caspian Terns (Hydroprogne caspia), a cosmopolitan, generalist piscivore, reared in captivity and fed ad libitum and restricted (ca. one‐third lower caloric intake) diets. Ad libitum‐fed chicks grew at similar rates and achieved a similar size at fledging as previously documented for chicks in the wild and had energetic demands that closely matched allometric predictions. We identified three general characteristics of food‐restricted Caspian Tern chicks compared to ad libitum chicks: (1) lower age‐specific body mass, (2) lower age‐specific skeletal and feather size, such as wing chord length, and (3) heightened levels of corticosterone in blood, both for baseline levels and in response to acute stress. Effects of diet restriction on feather growth (10–11% slower growth in diet‐restricted chicks) were less pronounced than effects on structural growth (37–52% slower growth) and body mass (24% lower at fledging age), apparently due to preferential allocation of food resources to maintain plumage growth. Our results suggest that measurements of chick body mass and feather development (e.g., wing chord or primary length) or measurement of corticosterone levels in the blood would allow useful evaluation of the nutritional status of chicks reared in the wild and of food availability in the foraging range of adults. Such evaluations could also inform demography studies (e.g., predict future recruitment) and assist in evaluating designated piscivorous waterbird conservation (colony) sites.  相似文献   

15.
In this study we examined hormonal responses of Black-legged Kittiwake (Rissatridactyla) chicks to experimental variations in energy content and nutritional quality (low or high lipid to protein ratio, LPR) of their food. Starting at the age of 10 days, chicks were fed either high or low LPR fish at 30, 50, 70 and 100% of ad libitum energy intake. After 20 days of treatment, chicks were exposed to a standardized acute handling and restraint stress protocol, where a baseline sample was taken immediately after taking a chick from the nest, and three additional blood samples were taken at intervals up to 50 min. Testosterone and corticosterone titres in plasma were measured via radioimmunoassay. We found that baseline testosterone levels were not significantly affected by the experimental treatments. Food-restricted chicks had elevated baseline and acute stress-induced levels of corticosterone compared to chicks fed ad libitum. An elevation of circulating levels of corticosterone in energetically stressed individuals was further magnified by low nutritional quality of food. Baseline and acute stress-induced corticosterone levels of chicks were negatively correlated with their fat reserves. We conclude that the physiological condition of Black-legged Kittiwake chicks can be assessed reliably by measuring circulating levels of corticosterone. We discuss short- and long-term effects of elevated corticosterone secretion in food-stressed nest-bound chicks. Accepted: 13 April 1999  相似文献   

16.
Abstract: Declining sage-grouse (Centrocercus urophasianus) populations may be characterized by poor recruitment largely attributed to low chick survival. However, few published studies have explicitly examined factors that influence chick survival. We used a suture method to radiomark 1-2-day-old sage-grouse chicks (n = 150) in 2005-2006 on Parker Mountain in south-central Utah, USA, and monitored their survival to 42 days. We modeled effects of year, hatch date, chick age, brood-female age, brood-mixing, and arthropod abundance on chick survival. Our best model revealed an average survival estimate of 0.50 days to 42 days, which is the highest level ever documented for this long-lived species. Brood-mixing occurred in 21% (31/146) of chicks and 43% (18/42) of broods we studied. Moreover, yearling females had more chicks leave their broods than did adults. We found that survival may be higher among chicks that switch broods compared to those that stayed with their natal mother until fledging. Thus, brood-mixing may be an adaptive strategy leading to increased sage-grouse chick survival and higher productivity, especially among chicks born to yearling females. Our findings also indicate that arthropod abundance may be an important driver of chick survival, particularly during the early brood-rearing period and, therefore, sage-grouse populations may benefit from a management strategy that attempts to increase arthropod abundance via brood habitat management.  相似文献   

17.
Bird nests are often heavily infested with several haematophagous ectoparasite species that drain energy and other essential resources needed for the development of the rapidly growing chicks. The nutritional requirements of altricial chicks can only be satisfied by the parents frequently bringing prey to the nest. In a 3-year study, we tested experimentally whether a parasite-induced change in the nutritional budgets of the chicks in an Algerian population of the Blue Tit, could be compensated by a response in parental food provisioning behaviour. We found that nestling body mass and size did not differ between heat-treated 'parasite-free' nests and heavily infested control nests. However, after controlling for potential confounding variables (laydate, clutch size, chick age, year), we found that broods of heavily infested control nests were more frequently visited and fed than broods that received antiparasite treatment. The results are discussed in the framework of theory related to behavioural responses of hosts to parasites.  相似文献   

18.
We present data on chick growth and chick feeding in Wilson's storm-petrel (Oceanites oceanicus) in a colony on King George Island, South Shetland Islands. Chicks were repeatedly weighed and the weight differences over 24 h were corrected for metabolic loss in order to obtain an estimation of meal sizes. Chicks were fed on 93% of the nights (n=688 nights). The average meal size for a single feeding was 8.5 g. Chicks received on average 1.2 feedings per night. These results are compared with data for this species from other locations. There was a trend for increased meal sizes from northern to southern populations, parallel to an increase in the adult mass, indicating that Wilson's storm-petrels carry optimal meal sizes according to their body size and may take advantage of increased food abundance by increasing feeding frequencies. We describe chick growth and discuss the influence of egg size, hatching date and feeding frequency on chick growth. The egg size had a positive influence on tarsus growth and body mass of chicks. Later-hatched chicks started wing growth and finished mass growth at a younger age and reached lower peak masses, indicating that late chicks may adapt to the restricted breeding season in their Antarctic breeding grounds by a more rapid development, but will fledge with a lower degree of development and less resources. Accepted: 22 May 2000  相似文献   

19.
Age and sex ratios in bag records are frequently used as indices of population composition for harvested populations. However, vulnerability to harvest may differ by age and sex thereby producing bias in population estimates. We assessed whether age and sex affected vulnerability to harvest for willow grouse (Lagopus lagopus) where adult density and brood size were known in the harvested populations. We collected bag records during 2 days of controlled hunting in 4 areas in 2 years (2007 and 2008) in Jämtland county, Sweden. We found that vulnerability to harvest was different for chicks and adults, but not between male and female adults. Hunters encountered broods at a higher rate than single birds compared to personnel conducting pre-harvest counts along line transects. Furthermore, the probability of shooting a grouse was higher in encounters of broods than individual grouse. Proportionally, we calculated about a 50% probability of a hunter shooting either a chick or an adult independent of encountering a single bird or broods of 2–10 grouse. Increasing adult density also increased the vulnerability to harvest for adults relative to chicks, independent of the chick to adult ratio in the pre-harvest population. The different vulnerability of adults and chicks to harvest observed in this study will dampen variation in age classes in bag records compared to the population, and we caution against extrapolation of age ratios in bag records to harvested populations. © 2012 The Wildlife Society.  相似文献   

20.
Grassland-breeding shorebirds show widespread declines due to a reduction in breeding productivity following agricultural intensification. However, there is also concern that increasing predation causes further declines or precludes population recovery. Predation may itself be enhanced by agriculture through changes in habitat or food availability, but little is known about the mortality of nidifugous shorebird chicks. We studied mortality by radio-tagging 662 chicks of Black-tailed Godwit Limosa limosa and Northern Lapwing Vanellus vanellus in 15 farmland sites in the Netherlands. Tagging and handling had no effect on the condition and survival of godwit chicks, but body condition was reduced by 6–11% in lapwing chicks wearing a tag for longer than 3 days. Fledging success was 0–24% in both species. Mortality was highest in young chicks but remained considerable until after fledging. Losses were traced mostly to predators (70–85%; 15 species, predominantly birds), but at least 5–10% were due to mowing, and 10–20% were due to other causes, including entrapment in ditches and starvation. Chicks staying in fields that were cut before the next radio check were found much more often as mowing victims and somewhat more often as prey remains than chicks in fields not cut, indicating that predation includes a limited amount of scavenging. The predation hazard for godwit chicks was higher in recently cut or grazed fields than in the tall, uncut grasslands they preferred, while that for lapwing chicks was lowest in grazed fields. In godwit chicks, poor body condition increased mortality risk, not only from starvation but also from other causes. Predation on godwit chicks was thus enhanced by intensive farming through a decline in the availability of cover, augmented by a reduced body condition, possibly due to food availability problems. Changes in farming practice may therefore help reduce predation pressure, though the observed interactions explained only part of the high predation rate in godwits and none in lapwings. Predator abundance has increased in Dutch wet grassland regions, and chick predation has become a factor that should be considered in planning the type and location of conservation measures.  相似文献   

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