Finch Eggs Research Articles

Effects of the temperature during embryonic development on adult reproduction and the phenotype of the second generation in zebra finches

Across taxa, the temperature experienced by individuals early in life can have large effects on their development. However, comparatively little is known about whether the effects of this thermal developmental environment can be long-lasting or transgenerational. In birds, one important aspect of the developmental environment is incubation and, in general, eggs incubated at low temperatures produce offspring with smaller morphology, suboptimal physiology, and even lower long-term survival. Yet, little is known about whether incubation temperature may affect avian reproduction in adulthood, and nothing is known about whether the effects of avian incubation temperature may be transgenerational. To investigate this, we incubated zebra finch (Taeniopygia guttata) eggs at two different temperatures: 37.5 °C (‘control’) and 36.3 °C (‘low’), raised nestlings until adulthood, and allowed same-temperature treatment pairs to reproduce. We found that F1 individuals incubated at the low temperature had shorter beaks at the start of reproduction than those incubated at the control temperature. Further, compared to those from control parents, F2 offspring from parents incubated at the low temperature had lighter body masses at 5 days-old and had shorter beaks at 30 days-old. However, we found little evidence that incubation temperature affected other aspects of reproduction, with no effect on latency to lay, clutch size, egg mass, incubation period, hatching success/asynchrony, fledging, or the number of offspring that ultimately survived until independence. Overall, we found some evidence that a difference in the early thermal developmental environment can have lasting morphological effects into the next generation. However, future work is needed to determine whether the incubation temperature that birds experience as embryos may influence parental care behaviors or lifetime reproductive success.

Repeatable randomness, invariant properties, and the design of biological signatures of identity.

What makes a perfect signature? Optimal signatures should be consistent within individuals and distinctive between individuals. In defense against avian brood parasitism, some host species have evolved "signatures" of identity on their eggs, comprising interindividual variation in color and pattern. Tawny-flanked prinia (Prinia subflava) egg signatures facilitate recognition and rejection of parasitic cuckoo finch (Anomalospiza imberbis) eggs. Here, we show that consistency and distinctiveness of patterns are negatively correlated in prinia eggs, perhaps because non-random, repeatable pattern generation mechanisms increase consistency but limit distinctiveness. We hypothesize that pattern properties which are repeatable within individuals but random between individuals ("invariant properties") allow hosts to circumvent this trade-off. To find invariant properties, we develop a method to quantify entire egg phenotypes from images taken from different perspectives. We find that marking scale (a fine-grained measure of size), but not marking orientation or position, is an invariant property in prinias. Hosts should therefore use differences in marking scale in egg recognition, but instead field experiments show that these differences do not predict rejection of conspecific eggs by prinias. Overall, we show that invariant properties allow consistency and distinctiveness to coexist, yet receiver behavior is not optimally tuned to make use of this information.

Combined measures of mimetic fidelity explain imperfect mimicry in a brood parasite-host system.

The persistence of imperfect mimicry in nature presents a challenge to mimicry theory. Some hypotheses for the existence of imperfect mimicry make differing predictions depending on how mimetic fidelity is measured. Here, we measure mimetic fidelity in a brood parasite-host system using both trait-based and response-based measures of mimetic fidelity. Cuckoo finches Anomalospiza imberbis lay imperfectly mimetic eggs that lack the fine scribbling characteristic of eggs of the tawny-flanked prinia Prinia subflava, a common host species. A trait-based discriminant analysis based on Minkowski functionals-that use geometric and topological morphometric methods related to egg pattern shape and coverage-reflects this consistent difference between host and parasite eggs. These methods could be applied to quantify other phenotypes including stripes and waved patterns. Furthermore, by painting scribbles onto cuckoo finch eggs and testing their rate of rejection compared to control eggs (i.e. a response-based approach to quantify mimetic fidelity), we show that prinias do not discriminate between eggs based on the absence of scribbles. Overall, our results support relaxed selection on cuckoo finches to mimic scribbles, since prinias do not respond differently to eggs with and without scribbles, despite the existence of this consistent trait difference.

Visual complexity of egg patterns predicts egg rejection according to Weber's law.

Visual complexity is ubiquitous in nature. Drivers of complexity include selection in coevolutionary arms races between antagonists. However, the causes and consequences of biological complexity and its perception are largely understudied, partly because complexity is difficult to quantify. Here, we address this by studying egg pattern complexity and its perception in hosts (tawny-flanked prinia Prinia subflava), which visually recognize and reject mimetic eggs of their virulent brood parasite (cuckoo finch Anomalospiza imberbis). Using field data and an optimization algorithm, we compute a complexity metric which predicts rejection of experimentally placed conspecific eggs in prinia nests. Real cuckoo finch eggs exhibit significantly lower pattern complexity than prinia eggs, suggesting that high complexity benefits hosts because it distinguishes host eggs from parasitic eggs. We show that prinias perceive complexity differences according to Weber's law of proportional processing (i.e. relative, rather than absolute, differences between stimuli are processed in discrimination, such that two eggs with simple patterns are more easily discriminable than two with complex patterns). This may influence coevolutionary trajectories of hosts and parasites. The new methods presented for quantifying complexity and its perception can help us to understand selection pressures driving the evolution of complexity and its consequences for species interactions.

Effects of prenatal testosterone on cumulative markers of oxidative damage to organs of young adult zebra finches (Taeniopygia guttata).

We tested the hypothesis that exposure of avian embryos to androgens in ovo entails long-term costs in the form of oxidative damage to vital cells and organs in adulthood. We injected zebra finch eggs with testosterone (T), monitored postnatal growth, and analyzed markers of oxidative damage in heart and liver in mature birds. We measured 8-oxo-2'-deoxyguanosine and isoprostanes, markers of oxidative damage to DNA and membrane lipids, respectively. T treatment (1) reduced growth rates of female but not male nestlings vs. controls; (2) resulted in less accumulation of 8-oxo-dG, but not IsoPs, in liver tissue of 60-day-old females, but not males; and (3) a trend toward elevated 8-oxo-dG levels in heart tissue of males and females at 60 and 180days old combined. These results generally support the testosterone oxidative damage hypothesis, in that embryonic exposure to higher T resulted in damage to DNA of heart tissue in both sexes. They also suggest that sex-specific effects of androgens on early growth rates may carry over as differences in some forms of oxidative damage in adults. This supports a basic tenet of evolutionary aging theory that developmental influences early in life can be linked to costs later on.

Experimental manipulation of maternal corticosterone: Hormone transfer to the yolk in the zebra finch Taeniopygia guttata

Maternally-derived hormones affect offspring physiological and behavioural phenotype, plausibly as an adaptive response to maternal environmental conditions. Corticosterone (CORT), the principal avian glucocorticoid produced in response to stress, is recognised as a potential mediator of such maternal reproductive effects. Maternally-derived yolk CORT is implicated in mediating offspring growth and hatchling begging behaviour. However, determining the potential for maternal effects in opportunistic breeders subject to variable environments relies on understanding whether natural variation in maternal circulating hormones may directly impact the embryo during development. Therefore, we tested whether elevated maternal CORT concentrations increase yolk CORT concentrations in zebra finch (Taeniopygia guttata) eggs. We remotely dosed breeding females with biologically-relevant doses of CORT, or the oil vehicle, 0–3 h prior to the predicted time of ovulation, and allowed pairs to produce two clutches, one under each treatment, in a crosswise, balanced design. CORT dosing elevated maternal plasma CORT and increased mean yolk CORT by a factor of 1.75 compared to the egg yolks of control mothers. Importantly, CORT concentrations did not differ between inner and outer layers of yolk. We found no egg lay order effect and maternal CORT dosing did not influence reproductive outputs (clutch initiation date, clutch size or egg mass). Our results confirm the direct impact of biologically-relevant increases in maternal CORT on yolk CORT, providing evidence that maternal CORT concentrations during yolk deposition to the follicle alters embryonic exogenous CORT exposure. Further research is required to determine the impact of maternal CORT on embryonic developmental programming.

Sex-Specific Effects of Incubation Temperature on Embryonic Development of Zebra Finch (Taeniopygia guttata) Embryos.

In oviparous species, the embryonic environment-particularly temperature-can alter phenotype and survival of an individual by affecting its size as well as its metabolic rate. Previous studies have shown that incubation temperatures can affect sex ratio in birds; specifically, low incubation temperatures were shown to produce a male-biased sex ratio in zebra finches (Taeniopygia guttata) possibly because of a higher pre- or postnatal mortality rate in females. We hypothesized that sexes respond differently to suboptimal incubation temperature, leading to a male-biased sex ratio. To test this hypothesis, zebra finch eggs were incubated at 36.1°, 37.5°, or 38.5°C and hatching success, hatchling mass, residual yolk mass, and pectoralis mass were measured. We found that while hatchling mass was similar between the sexes at 37.5°C, female hatchlings were heavier at 36.1°C, and male hatchlings were heavier at 38.5°C. Pectoralis muscle mass was similar between the sexes at 36.1°C; however, at 37.5°C, female pectoralis mass was heavier at hatching than that of males. Females at 37.5°C also had lower residual yolk at hatching compared with males, reflecting a higher use of energy by female embryos compared with male embryos at this temperature. In contrast, residual yolk was similar between the sexes at 36.1° and 38.5°C. Our results suggest that there are sex differences in how incubation temperature alters organ mass and yolk energy reserve; this can lead to a difference in survival at different incubation temperatures between the sexes. Taken together with previous studies showing that females alter incubation behavior with ambient temperature, rising ambient temperatures could impact phenotype and survival of avian offspring in a sex-specific manner.

In ovo tris(2-butoxyethyl) phosphate concentrations significantly decrease in late incubation after a single exposure via injection, with no evidence of effects on hatching success or latent effects on growth or reproduction in zebra finches.

Zebra finch (Taeniopygia guttata) eggs were injected with the organophosphate triester flame retardant tris(2-butoxyethyl) phosphate (TBOEP) at 0 μg/g, 0.01 μg/g, 1 μg/g, 10 μg/g, or 50 μg/g egg. Subsets of high-dose eggs were collected throughout incubation to measure TBOEP, which started declining in late incubation and then decreased rapidly to 28% of injected concentration by hatching. The authors found no effects of TBOEP on survival, growth, or reproduction even at very high doses. Environ Toxicol Chem 2017;36:83-88. © 2016 SETAC.

Zebra finch as a developmental model.

The domesticated zebra finch (Taeniopygia guttata) is a well-established animal model for studying vocal learning. It is also a tractable model for developmental analyses. The finch genome has been sequenced and methods for its transgenesis have been reported. Hatching and sexual maturation in this species takes only two weeks and three months, respectively. Finch colonies can be established relatively easily and its eggs are laid at a stage earlier than in other common avian experimental models, facilitating the analysis of very early avian development. Representing the Neoaves to which 95% of all bird species belong, the finch can potentially complement two existing, Galloanserae developmental models, the chick, and quail. Here, we provide a step-by-step guide for how to set up a finch colony in a conventional laboratory environment. Technical tips are offered to optimize hens' productivity and ensure a constant supply of fertilized finch eggs. Methods of handling finch eggs and embryos for subsequent embryological, cellular, or molecular analyses are also discussed. We conclude by emphasizing scientific values and cost effectiveness of maintaining a finch colony for avian developmental studies. genesis 53:669-677, 2015. © 2015 Wiley Periodicals, Inc.

Host-Parasite Arms Races and Rapid Changes in Bird Egg Appearance

Coevolutionary arms races are a powerful force driving evolution, adaptation, and diversification. They can generate phenotypic polymorphisms that render it harder for a coevolving parasite or predator to exploit any one individual of a given species. In birds, egg polymorphisms should be an effective defense against mimetic brood parasites and are extreme in the African tawny-flanked prinia (Prinia subflava) and its parasite, the cuckoo finch (Anomalospiza imberbis). Here we use models of avian visual perception to analyze the appearance of prinia and cuckoo finch eggs from the same location over 40 years. We show that the two interacting populations have experienced rapid changes in egg traits. Egg colors of both species have diversified over time, expanding into avian color space as expected under negative frequency-dependent selection. Egg pattern showed signatures of both frequency-dependent and directional selection in different traits, which appeared to be evolving independently of one another. Host and parasite appear to be closely tracking one another's evolution, since parasites showed closer color mimicry of contemporaneous hosts. This correlational evidence suggests that hosts and parasites are locked in an ongoing arms race in egg appearance, driven by constant change in the selective advantage of different phenotypes, and that coevolutionary arms races can generate remarkably rapid phenotypic change.

Tools for managing invasions: acceptance of non-toxic baits by juvenile Nile monitor lizards and Burmese pythons under laboratory conditions

Nile monitor lizards (Varanus niloticus) and Burmese pythons (Python molurus bivittatus) are large, invasive, predatory reptiles, which are now well established in south Florida. Acetaminophen was recently shown to be lethal to both animals and therefore has potential for inclusion in an integrated pest management effort to control these species. However, acceptable bait matrices for both species are still needed to deliver the toxicant. We tested nine candidate bait matrices on juvenile Nile monitor lizards and Burmese pythons in the laboratory. Baits were tested fresh and also after aging at 30°C and 50% relative humidity for 24 h. Six of the nine baits were well accepted by the monitors in both fresh and aged conditions (dead neonatal mouse, dead quail chick, ground turkey, chicken liver, tilapia, and zebra finch egg). Only the fresh dead neonatal mouse and fresh dead quail chick were well accepted by the pythons. As far as we are aware, these tests are the second of such bait matrix preference tests conducted for reptiles. The implications of our results are discussed regarding further development and testing of baits in a natural setting.

Yolk androgens and embryo sex: Maternal effects or confounding factors?

Maternal effects occur when offspring phenotype is affected by environmental factors experienced by the mother and, in egg-laying species, are often mediated via egg resources. There is currently great interest among behavioural ecologists in maternally allocated yolk androgens, especially their relationship with offspring sex and development. Such studies need embryonic tissue for sexing, however, requiring eggs to be incubated (usually for 3 days). Therefore, there are concerns about whether the androgen concentrations assayed reflect those allocated by the mother. In addition, studies showing sex biases in maternal allocation of androgens could be confounded if male and female embryos uptake or metabolise androgens at different rates. We ran a series of experiments using zebra finch ( Taeniopygia guttata) eggs to address these potential confounding factors. First we showed, using eggs naturally incubated for up to 5 days, that eggs containing embryos had lower yolk androgen concentrations than eggs that had failed to form embryos. We then tested various hypotheses for this difference using controlled incubation treatments. Our results suggested that (a) embryo development causes the yolk to become progressively more diluted with albumin; and (b) between 3 and 5 days of incubation embryos start uptaking or metabolising androgens. Crucially, we found no decline in yolk androgen concentration at 3 days incubation, and no evidence for sex-specific rates of uptake or metabolism of androgens. This strongly suggests that yolk androgen levels up to 3 days incubation do reflect those allocated by the mother, and that studies of sex biased maternal allocation of yolk androgens are not confounded by sex differences in embryo development.

Carolina Wren Nest Successfully Parasitized by House Finch

We report the first observation of a House Finch (Carpodacus mexicanus) successfully parasitizing a Carolina Wren (Thryothorus ludovicianus) nest. On 24 May 2005, we found a Carolina Wren nest in south-central Oklahoma containing four Carolina Wren eggs and two House Finch eggs. The House Finch eggs hatched and nestlings grew rapidly. The Carolina Wren eggs hatched but the young did not survive. We observed a House Finch fledgling with the adult Carolina Wrens the day after fledging.

Yolk Testosterone Stimulates Growth and Immunity in House Finch Chicks

Female birds deposit variable amounts of androgens, such as testosterone, into the yolks of their eggs. Evidence suggests that yolk androgens play an important role in the determination of offspring phenotype. While androgens are generally regarded as anabolic and immunosuppressive, studies of the behavioral and physiological effects of yolk androgens on offspring of several avian species have been conflicting, leaving the adaptive significance associated with deposition patterns of yolk androgens unclear. We injected either a physiological dose of testosterone or a control vehicle into house finch (Carpodacus mexicanus) eggs and examined the effects of these injections on offspring growth and immunity. Two days after hatching, nestlings from eggs treated with testosterone were significantly larger than nestlings from eggs treated with a control injection, suggesting a stimulatory effect of yolk androgens in early development. By 8 d after hatching, however, this effect disappeared, and chicks from the two treatment groups were similar in size. Nestlings in the testosterone treatment group showed a significantly larger swelling response to phytohemagglutinin than control nestlings 15 d after hatching, which is close to fledging. Overall, our observations show that when food resources are abundant, testosterone stimulates both early growth and immunity in developing house finches.

Sex allocation in response to paternal attractiveness in the zebra finch

Females mated to attractive males are predicted to produce male-biased broods. Previous studies on zebra finches, Taeniopygia guttata, in which colored leg rings were used to alter male attractiveness, support this hypothesis. However, because molecular sexing techniques were not available, it was not known when during development this bias arose. Also, because both attractive (red-ringed) and unattractive (green-ringed) males were within the same aviary, assortative mating between treatments may have confounded the results. Using two different experimental designs, we tested whether the sex ratio of zebra finch eggs and chicks differed in response to paternal ring color whilst controlling for assortative mating between treatments. In the aviary experiment, birds could interact socially, but all males in an aviary had the same leg ring color. In the cage experiment, each female was randomly assigned a red- or green-ringed mate, thus also eliminating assortative mating within treatments. Offspring were sexed based on plumage or using a molecular method. The sex ratio at laying did not differ between treatments in either the aviary (n = 313 eggs) or cage (n = 151 eggs) experiments, suggesting that female zebra finches do not manipulate the primary sex ratio in response to their mate's ring color. However, in the cage experiment we found greater male embryonic mortality in the attractive group, which resulted in a female-biased sex ratio at sexual maturity, that is, in the opposite direction to that found in previous studies. Possible explanations for the disparity between our results and those of previous studies are considered.

Sex-biased investment in yolk androgens depends on female quality and laying order in zebra finches (Taeniopygia guttata)

The Trivers-Willard hypothesis predicts sex biases in parental investment according to parental condition. In addition, parents may need to sex bias their investment if there is an asymmetry between the sexes in offspring fitness under different conditions. For studying maternal differential investment, egg resources are ideal subjects because they are self contained and allocated unequivocally by the female. Recent studies show that yolk androgens can be beneficial to offspring, so here we test for sex-biased investment with maternal investment of yolk testosterone (T) in zebra finch (Taeniopygia guttata) eggs. From the Trivers-Willard hypothesis, we predicted females to invest more in male eggs in optimum circumstances (e.g. good-condition mother, early-laid egg), and more in female eggs under suboptimal conditions (e.g. poor-condition mother, late-laid egg). This latter prediction is also because in this species there is a female nestling disadvantage in poor conditions and we expected mothers to help compensate for this in female eggs. Indeed, we found more yolk T in female than male eggs. Moreover, in accordance with our predictions, yolk T in male eggs increased with maternal quality relative to female eggs, and decreased with laying order relative to female eggs. This supports our predictions for the different needs and value of male and female offspring in zebra finches. Our results support the idea that females may use yolk androgens as a tool to adaptively manipulate the inequalities between different nestlings.

Subtle manipulation of egg sex ratio in birds

Mothers are predicted to overproduce male or female eggs when the relative fitness gains from one sex are higher and outweigh the costs of manipulation. However, in birds such biases are often difficult to distinguish from differential embryo or chick mortality. Using a molecular technique to identify the sex of early embryos, we aim to determine the effect of maternal nutrition on zebra finch (Taeniopygia guttata) egg sex ratios after 2 days of incubation, which is as close to conception as is currently possible. We found no overall bias in the sex ratio of eggs laid and sex did not differ with relative laying order under any diet regime. However, mothers on a low-quality diet did produce a female bias in small clutches and a slight male bias in large clutches. On a high-quality diet, mothers produced a male bias in small clutches and a female bias in large clutches. Those on a standard diet produced a roughly even sex ratio, irrespective of clutch size. These observed biases in egg sex are partly in line with predictions that, in this species, daughters suffer disproportionately from poor rearing conditions. Thus, when relatively malnourished, mothers should only rear daughters in small broods and vice versa. Sex-ratio patterns in this species therefore appear to be subtle.

The effect of variation in dietary intake on maternal deposition of antioxidants in zebra finch eggs

Summary Maternal diet can significantly influence the quality and size of eggs, and this may, in turn, influence the fitness of offspring. In this study, we show how antioxidants (vitamin E compounds and carotenoids) in the diet of female zebra finches influence the concentrations of antioxidants in their eggs. Antioxidants are biochemicals derived from the diet, which reduce damage to cell membranes caused by the free radicals produced during normal metabolism and growth. Females were given either a seed‐only or a seed + rearing food diet prior to egg laying. The seed + rearing food diet was more enriched with carotenoids, but had lower amounts of less effective antioxidants, such as γ‐tocopherol. Eggs were collected as they were laid, then analysed to determine the concentrations of antioxidants using high‐performance liquid chromatography. Females in the two food groups had similar rates of food consumption and laid similar sized clutches. Females on the seed‐only diet produced eggs of similar mass to that of females on the seed + rearing food diet. The concentrations of the most effective antioxidant (α‐tocopherol) were higher in the eggs of females from the seed‐only group and, for both food groups, concentrations decreased with successively laid eggs within clutches. The concentration of carotenoids in egg yolks did not differ between food treatments, but also decreased with successive eggs. Less effective antioxidants were relatively under‐represented in eggs in relation to their availability in food, and were deposited in similar overall amounts between the two food treatments, and independently of laying sequence. Our results indicate that egg size is not necessarily an accurate guide to egg quality, especially when comparing between clutches of different females or across species. Females may be able to adjust egg composition to influence offspring quality variation within broods, possibly traded‐off against their personal use of antioxidants, their future reproductive success, control over sibling competition within their brood and/or brood reduction.

Predator Activity and Nest Success of Willow Flycatchers and Yellow Warblers

Willow flycatchers (Empidonax traillii) and yellow warblers (Dendroica petechia) are riparian-dependent species that have declined throughout much of their former range in California, USA. These declines have been primarily associated with the loss of riparian breeding habitat, increases in brood parasitism, and increases in nest predation. We (1) identified potential nest predators using inactive yellow warbler nests; (2) determined the relationship of meadow wetness, meadow size, and amount of edge to predator activity; (3) determined the association between potential nest predator activity and nest success; and (4) determined how proximity to forest edge and isolated trees was related to nest success. We used automatic cameras to monitor inactive yellow warbler nests baited with zebra finch (Taeniopygia guttata) eggs to identify nest predators. We used track plates (mammalian), point counts (avian), and time-constrained searches (reptilian) to assess the activity of potential nest predators. We photographed short-tailed weasel (Mustela erminea), Douglas squirrel (Tamiasciurus douglasii), lodgepole chipmunk (Tamias speciosus), deer mouse (Peromyscus maniculatus), and unidentified chipmunks (Tamias spp.) depredating yellow warbler nests baited with finch eggs. The amount of meadow covered with water was negatively associated with the activity of chipmunks and Douglas squirrels. Meadow size was negatively associated with Douglas squirrel activity. The amount of edge was positively associated with the activity of Douglas squirrels, chipmunks, Steller's jays (Cyanocitta stelleri), and brown-headed cowbirds (Molothrus ater). Nest predation was the major cause of nest failure in our study. However, only short-tailed weasels, Douglas squirrels, Clark's nutcrackers (Nucifraga columbiana), Steller's jays, Cooper's hawks (Accipiter cooperii), and brown-headed cowbirds had activity indices that were negatively associated with nest success of either species. The distance to isolated trees was associated with willow flycatcher nest success, whereas the distance to both isolated trees and to the forest edge was associated with yellow warbler nest success-nests located closer to isolated trees and the forest edge were more likely to be parasitized and/or depredated. Our results suggest that flooding portions of meadows may restrict meadow access to forest-edge-associated nest predators.

Measuring Edge Effects on Nest Predation in Forest Fragments: Do Finch and Quail Eggs Tell Different Stories?

Experiments assessing rates of avian nest predation often find that nests near forest edges are at high risk of predation, suggesting the importance of forest fragmentation in recent population declines of ground-nesting passerines. However, the use of quail (Coturnix spp.) eggs in nest predation experiments may confound conclusions about edge effects because only large-mouthed predators are able to consume these relatively large eggs, but both large and small-mouthed predators consume smaller passerine eggs. We directly compared predation rates on artificial nests baited with quail eggs or with zebra finch (Poephila guttata) eggs; the latter are similar in size to the eggs of many neotropical passerines. In 1998 and 1999 we placed 392 artificial ground nests at edge and interior locations in two east-central Iowa forest fragments. Predation on these nests varied with egg type (quail or finch) and location (edge or interior) and there was a significant interaction between egg type and location: predation on quail eggs was greater at edges than in the interior, whereas finch egg predation was high in both edge and interior locations. Based on tooth imprints in clay eggs, we determined that large-mouthed predators were six times more active at edges, whereas activity of small-mouthed nest predators was evenly distributed between edge and interior locations. We suggest that the use of only quail eggs can exaggerate edge effects and that finch eggs or clay eggs used in conjunction with quail eggs in artificial nests can be used to estimate relative predation rates by large- and small-mouthed predators.