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USGS1974

The literature of the California black rail

Few birds have remained so little known as the California black rail ( Laterallus jamaicensis coturniculus ). Although first collected in 1859 or before and reported in 1874 (Ridgway 1874), its life history, distribution, and status have remained so obscure that even a sight record of the bird is deemed worthy of a report in some ornithological publications. Because degradation and loss of freshwater and saltwater marshlands in California may be detrimentally affecting the black rail, both the U.S. Bureau of Sport Fisheries and Wildlife (1973) and the California Department of Fish and Game (1972) have classified it as rare and worthy of further study. The 84 papers and notes both summarized in this report and included in its bibliography include essentially all that is currently known about the California black rail. Only 11 of these papers consider the life history of this rail in any detail. The rest are distribution notes and some of the more important papers on the closely related eastern black rail ( L. j. jamaicensis ). The latter are included for comparative purposes, or because they may lend clues to currently unknown facets of the life history of the western race.

Special Scientific Report - Wildlife↗

USGS2007

The 1999-2003 Summary of the North American Breeding Bird Survey

Data from the North American Breeding Bird Survey were used to estimate continental and regional changes in bird populations for the 5-yr period 1999-2003 and the 2-yr period 2002-2003. These short-term changes were placed in the context of population trends estimated over the 1966-2003 interval. During 1999-2003, 41% of all species exhibited positive trends over the entire survey area, while 64% of all species exhibited positive change between 2002-2003. The continental and regional percentages of species with positive trends were also analyzed for 12 species groups having shared life-history traits. Survey-wide for the entire survey period, grassland birds exhibited the lowest percentage of increasing species (14%), with their sharpest declines occurring in the West during 1999-2003 (10% increasing). During 1999-2003, short-distance migrants experienced significant declines in all regions, where numbers of species with increasing trends ranged from 22% - 34%. Most species fared well during the 2002-2003 period, with 64% (P < 0.05) increasing survey-wide. This was primarily a result of increases in the Central and Western BBS regions where 21 of 24 species groups exhibited significant increases in the number of species with positive trends.

Bird Populations↗

USGS2001

Reproductive ecology and demography of the 'Akohekohe

The ‘Ākohekohe ( Palmeria dolei ) is an endangered Hawaiian honeycreeper endemic to the montane rain forests of east Maui in the Hawaiian Islands. We investigated ‘Ākohekohe nesting ecology using color-banded birds for the first time as a background to understanding the species' conservation. From 1994–1997, we color-banded 78 individuals, located and monitored 46 active nests, and took behavioral data during 534 hr of nest observation at Hanawī Natural Area Reserve, near the center of the species' range. ‘Ākohekohe nesting behavior and life history closely resembled that of ‘Apapane ( Himatione sanguinea ) and related honeycreepers. The birds were monogamous within and among years, and we found no evidence of polyandry, polygyny, or helpers at the nest. The nesting season extended from November to early June. Females performed all incubation and brooding. Males provisioned females and nestlings, and they were more active than females in feeding fledglings during the two-week period of parental dependency. Modal clutch size, as determined from egg counts at three nests and by counting begging chicks at other nests, was two eggs, and parents frequently fledged two chicks. We found an overall nest success rate of 68% by the Mayfield method, a high rate compared with other Hawaiian honeycreepers and continental passerines. An average of 1.1 chicks fledged per active nest, and at least 42% of nesting pairs made two or more nesting attempts per season. Rats ( Rattus spp.) were abundant at the study site, and we confirmed their depredating some ‘Ākohekohe nests, so we did not expect to find such a high rate of nest success. The estimated annual probability of adult survival was also high, at 0.95 ± 0.10 (SE).

Condor↗

USGS2000

The 1995-1999 Summary of the North American Breeding Bird Survey

Data from the North American Breeding Bird Survey were used to estimate continental and regional changes in bird populations for the 5-yr period 1995-1999 and the 2-yr period 1998-1999. These short-term changes were placed in the context of population trends estimated over the 1966-1999 interval. During 1995-1999, 44% of all species exhibited positive trends over the entire survey area, while 44% of all species exhibited positive trends during 1998-1999; neither of these percentages differed significantly from 50%. The continental and regional percentages of species with positive trends were also analyzed for 12 species groups having shared life-history traits. Survey-wide for the entire survey period, grassland birds exhibited the lowest percentage of increasing species (19%). However, during 1995-1999 the declines were less extreme in the Central and Western BBS regions, with 49% and 36% of species increasing in these regions. Neotropical migrants continued to fare better than grassland birds in all regions, although in the Eastern BBS region only 30% of neotropical species had increasing trends during 1995-1999.

Bird Populations↗

USGS2011

Microanatomy of passerine hard-cornified tissues: Beak and claw structure of the black-capped chickadee (Poecile atricapillus)

The microanatomy of healthy beaks and claws in passerine birds has not been well described in the literature, despite the importance of these structures in avian life. Histological processing of hard‐cornified tissues is notoriously challenging and only a few reports on effective techniques have been published. An emerging epizootic of beak deformities among wild birds in Alaska and the Pacific Northwest region of North America recently highlighted the need for additional baseline information about avian hard‐cornified structures. In this study, we examine the beak and claw of the Black‐capped Chickadee ( Poecile atricapillus ), a common North American passerine that is affected by what has been described as “avian keratin disorder.” We use light and scanning electron microscopy and high‐magnification radiography to document the healthy microanatomy of these tissues and identify features of functional importance. We also describe detailed methods for histological processing of avian hard‐cornified structures and discuss the utility of special stains. Results from this study will assist in future research on the functional anatomy and pathology of hard‐cornified structures and will provide a necessary reference for ongoing investigations of avian keratin disorder in Black‐capped Chickadees and other wild passerine species.

Journal of Morphology↗

USGS2020

Determinants and consequences of dispersal in vertebrates with complex life cycles: a review of pond-breeding amphibians

Dispersal is a central process in ecology and evolution. It strongly influences the dynamics of spatially structured populations, by affecting population growth rate and local colonization-extinction processes. Dispersal can also influence evolutionary processes because it determines rates and patterns of gene flow in spatially structured populations and is closely linked to local adaptation. For these reasons, dispersal has received considerable attention from ecologists and evolutionary biologists. However, although it has been studied extensively in taxa such as birds and mammals, much less is known about dispersal in vertebrates with complex life cycles such as pond-breeding amphibians. Over the past two decades, researchers have taken an interest in amphibian dispersal and initiated both fundamental and applied studies, using a broad range of experimental and observational approaches. This body of research reveals complex dispersal patterns, causations and syndromes, with dramatic consequences for the demography and genetics of amphibian populations. In this review, our goals are to (1) redefine and clarify the concept of amphibian dispersal, (2) review current knowledge about the effects of individual (i.e., condition-dependent dispersal) and environmental (i.e., context-dependent dispersal) factors during the three stages of dispersal (i.e., emigration, immigration, transience), (3) identify the demographic and genetic consequences of dispersal in spatially structured amphibian populations, and (4) propose new research avenues to extend our understanding of amphibian dispersal. In particular, we emphasize the need to (1) quantify dispersal rate and distance rigorously using suitable model systems, (2) investigate the genetic basis and dispersal evolution patterns, and (3) examine dispersal-related eco-evolutionary dynamics. These proposed research avenues tap from the recent advances in quantitative and molecular methods and have the potential to improve our understanding of dispersal in organisms with complex life cycles.

The Quarterly Review of Biology↗

USGS1989

Pulmonary lesions in disseminated visceral coccidiosis of sandhill and whooping cranes

Fifty cranes, consisting of 46 sandhill (Grus canadensis) and four whooping cranes (Grus americana), were studied. Eighteen sandhill cranes and the four whooping cranes were naturally infected with disseminated visceral coccidiosis (DVC). The remaining sandhill cranes were chicks experimentally infected with oocysts of Eimeria reichenowi and/or E. gruis; five chicks served as controls. There were no clinical signs attributed to respiratory infection. Necropsy of naturally infected adult birds revealed nodules in many organs, including the lung, air sacs, trachea and nares. Artificially infected sandhill cranes and the whooping crane chicks that died from DVC had congestion and consolidated areas in the lung with frothy fluid in the airways. Grossly visible nodules were observed from 10 days postinoculation. Granulomatous pneumonia and tracheitis were observed with light microscopy. Lesions were associated with merogonic and gametogonic stages of eimerian coccidia. Granulomas and granulomatous foci contained parasitized large mononuclear cells. Merogonic stages were seen in lymphoid cells by ultrastructural examination. Oocysts were observed in the trachea and bronchial mucosa and admixed with exudate in the airways, indicating that crane eimerians can complete their life cycle at these sites. Of the few eimeriid coccidia that have extraintestinal stages of development in birds and mammals, only the species in cranes complete their life cycle in both the digestive and respiratory tracts.

Journal of Wildlife Diseases↗

USGS2024

North American bird banding program longevity records

Longevity in wild birds refers to the longest-lived individual of any particular species. This measure of age can reflect important life history traits and identifying differences in longevity patterns between species can reveal unique threats or conservation needs. Because it requires the identification of individual birds, longevity can usually only be measured through banding or marking of individuals. A bird’s age can generally be determined at the time of banding and then recalculated whenever the bird is subsequently recovered, recaptured, or resighted. The North American Bird Banding Program (NABBP), comprised of the U.S. Geological Survey Eastern Ecological Science Center Bird Banding Laboratory (BBL) and Canada’s Bird Banding Office (BBO), curates a vast dataset of all North American bird banding records, as well as subsequent encounters of banded birds reported by banders and members of the public. Although not all bird populations are banded equally and band encounters are only reported voluntarily, the NABBP is thus uniquely situated to catalogue and report longevity records for bird species under our purview.

North American Bird Bander↗

USGS2010

The 2003-2008 summary of the North American Breeding Bird Survey

Data from the North American Breeding Bird Survey were used to estimate continental and regional changes in bird populations for the 6-yr period 2003-2008 and the 2-yr period 2007-2008. These short-term changes were placed in the context of population trends estimated over the 1966-2008 interval. Across the entire survey area, a higher proportion of species exhibited positive growth during 2003-2008 (64%) than during the long-term (46%) or the more recent 2-yr-term (39%). The 2003-2008 growth occurred relatively evenly across the Western, Central, and Eastern BBS regions, with 59%, 66%, and 61% of all species increasing, respectively. We additionally evaluated the proportion of species with positive trend estimates in each of 12 life-history based groupings at continental and regional levels. Survey-wide, birds in the grassland guild demonstrated the lowest proportion of positive trends over the entire survey period (21% increasing), with significant declines occurring in both the Eastern and Western regions (5% increasing and 18% increasing, respectively). Birds in the wetland breeding guild exhibited the greatest proportion of positive trends, with a significant number of increasing species (between 77-90%) occurring in all three BBS regions during 2003-2008.

Bird Populations↗

USGS2014

Multiple plumage traits convey information about age and within-age-class qualities of a canopy-dwelling songbird, the Cerulean Warbler

Colorful plumage traits in birds may convey multiple, redundant, or unreliable messages about an individual. Plumage may reliably convey information about disparate qualities such as age, condition, and parental ability because discrete tracts of feathers may cause individuals to incur different intrinsic or extrinsic costs. Few studies have examined the information content of plumage in a species that inhabits forest canopies, a habitat with unique light environments and selective pressures. We investigated the information content of four plumage patches (blue-green crown and rump, tail white, and black breast band) in a canopy-dwelling species, the Cerulean Warbler ( Setophaga cerulea ), in relation to age, condition, provisioning, and reproduction. We found that older males displayed wider breast bands, greater tail white, and crown and rump feathers with greater blue-green (435–534 nm) chroma and hue than males in their first potential breeding season. In turn, older birds were in better condition (short and long term) and were reproductively superior to younger birds. We propose that these age-related plumage differences (i.e. delayed plumage maturation) were not a consequence of a life history strategy but instead resulted from constraints during early feather molts. Within age classes, we found evidence to support the multiple messages hypothesis. Birds with greater tail white molted tails in faster, those with more exaggerated rump plumage (lower hue, greater blue-green chroma) provisioned more, and those with lower rump blue-green chroma were in better condition. Despite evidence of reliable signaling in this species, we found no strong relationships between plumage and reproductive performance, potentially because factors other than individual differences more strongly influenced fecundity.

The Auk↗

USGS2024

Return rates of GPS-tagged Pacific Golden-Plovers: A controlled study in Hawaii

In a study of GPS-tagged Pacific Golden-Plovers wintering at Moorea, French Polynesia, Johnson et al. (2020) raised questions about possible tag-induced mortality. Similar concerns in other species have generated considerable attention in recent years. Of 19 tagged plovers that departed Moorea on northward migration, only one bird (5 %) uploaded a complete migratory cycle to Argos. Furthermore, contact was lost with 12 birds (63 %) variously during overwater flight, at stopover sites, or on nesting grounds; and no signals at all were received from 6 individuals (32%). Such observations suggest that carrying a tag might be a life-threatening burden. We cannot exclude the possibility that tag weight caused difficulties for the Moorea birds. However, this seems an unlikely factor in that the device (tag plus harness) being carried weighed less than 3 % of body mass (see beyond). Such a load is in accord with generally accepted albeit arbitrary limits. Other tag-associated concerns such as altered aerodynamics, increased vulnerability to predation, and (given the complex nature of geomagnetic navigation, the possibility that tag electronics might interfere with sense of direction, are more difficult to evaluate. Alternatively, loss of contact with tagged birds may be simply a matter of tag malfunction, battery failure, or a broken harness. To shed light on these uncertainties, and to explore the survival of GPS-tagged plovers on a nonbreeding ground in Hawaii, we conducted a controlled study using birds that winter in the National Memorial Cemetery of the Pacific (known locally as the Punchbowl).

Hawaii↗

USGS2023

Wetland management practices and secretive marsh bird habitat in the Mississippi Flyway: A review

Management regimes on publicly owned freshwater wetlands in the Mississippi Flyway of North America (i.e., Flyway) have historically emphasized waterfowl, but there is limited information on how waterfowl-focused wetland management affects other wetland-dependent wildlife. Secretive marsh birds (SMBs) depend on wetlands with emergent vegetation throughout their migratory life cycle and often encounter vegetation and water conditions resulting from waterfowl-focused management regimes. Thus, there is a need for better understanding of how SMBs are affected by wetland management and the extent to which waterfowl-focused management regimes provide habitat for SMBs. In this review, we identify the vegetation and water conditions resulting from typical management objectives on freshwater emergent wetlands in the Flyway, review and qualitatively synthesize results from studies that directly evaluate how wetland management practices affect SMBs or their habitat, and assess how the vegetation and water conditions being produced for target species (mainly waterfowl) align with SMB habitat requirements. We searched online databases and used Google Scholar to locate peer-reviewed literature, technical reports, and graduate theses that pertained to responses of SMBs or their habitat to water-level manipulation, herbicide application, prescribed fire, disking, mowing, and planting crops. There are several management strategies that complement SMBs and waterfowl, such as reducing cover of woody species and providing flooded emergent vegetation. We also highlight management strategies that may not currently align with SMB life-cycle needs and suggest adjustments that might promote habitat for SMBs while still achieving waterfowl population objectives. For example, adjusting the dates and duration of spring water-level drawdowns on a portion of wetlands within a larger complex can provide for spring migrating waterfowl and ensure habitat for migrating and nesting SMBs. Ideally, future studies would address how modifications to management practices affect SMBs and monitor potential effects on waterfowl, resulting in a more holistic approach to wetland management.

The Journal of Wildlife Management↗

USGS2026

Critter Tales—Adventures in restoration activity book for kids

Product Information “Critter Tales: Adventures in Restoration Activity Book” is an interactive educational resource designed for children ages 5–9, produced by the U.S. Geological Survey in cooperation with the U.S. Department of the Interior, Office of Restoration and Damage Assessment. Through the journey of Lulu the loon and a cast of animal friends, young readers explore the importance of ecological restoration across diverse habitats—from sand dunes and wetlands to prairies and streams. The activity book blends storytelling, science, and hands-on activities to teach children how restoration efforts help wildlife thrive. Readers learn about native plants, animal life cycles, and the role of restoration in protecting habitats for birds, turtles, butterflies, mussels, and more. Engaging puzzles, coloring pages, and interactive challenges encourage kids to become nature’s helpers, fostering environmental stewardship and awareness. With vibrant illustrations and accessible language, the publication highlights practical actions—such as planting native milkweed, using lead-free fishing tackle, and supporting mussel restoration—that empower children to make a positive change in their own communities. The book also introduces basic ecological concepts and celebrates the interconnectedness of living things, making it an ideal resource for families, educators, and anyone interested in environmental education.

General Information Product↗

USGS2010

Biological pathways of exposure and ecotoxicity values for uranium and associated radionuclides: Chapter D in Hydrological, geological, and biological site characterization of breccia pipe uranium deposits in Northern Arizona

This chapter compiles available chemical and radiation toxicity information for plants and animals from the scientific literature on naturally occurring uranium and associated radionuclides. Specifically, chemical and radiation hazards associated with radionuclides in the uranium decay series including uranium, thallium, thorium, bismuth, radium, radon, protactinium, polonium, actinium, and francium were the focus of the literature compilation. In addition, exposure pathways and a food web specific to the segregation areas were developed. Major biological exposure pathways considered were ingestion, inhalation, absorption, and bioaccumulation, and biota categories included microbes, invertebrates, plants, fishes, amphibians, reptiles, birds, and mammals. These data were developed for incorporation into a risk assessment to be conducted as part of an environmental impact statement for the Bureau of Land Management, which would identify representative plants and animals and their relative sensitivities to exposure of uranium and associated radionuclides. This chapter provides pertinent information to aid in the development of such an ecological risk assessment but does not estimate or derive guidance thresholds for radionuclides associated with uranium. Previous studies have not attempted to quantify the risks to biota caused directly by the chemical or radiation releases at uranium mining sites, although some information is available for uranium mill tailings and uranium mine closure activities. Research into the biological impacts of uranium exposure is strongly biased towards human health and exposure related to enriched or depleted uranium associated with the nuclear energy industry rather than naturally occurring uranium associated with uranium mining. Nevertheless, studies have reported that uranium and other radionuclides can affect the survival, growth, and reproduction of plants and animals. Exposure to chemical and radiation hazards is influenced by a plant’s or an animal’s life history and surrounding environment. Various species of plants, invertebrates, fishes, amphibians, reptiles, birds, and mammals found in the segregation areas that are considered species of concern by State and Federal agencies were included in the development of the site-specific food web. The utilization of subterranean habitats (burrows in uranium-rich areas, burrows in waste rock piles or reclaimed mining areas, mine tunnels) in the seasonally variable but consistently hot, arid environment is of particular concern in the segregation areas. Certain species of reptiles, amphibians, birds, and mammals in the segregation areas spend significant amounts of time in burrows where they can inhale or ingest uranium and other radionuclides through digging, eating, preening, and hibernating. Herbivores may also be exposed though the ingestion of radionuclides that have been aerially deposited on vegetation. Measured tissues concentrations of uranium and other radionuclides are not available for any species of concern in the segregation areas. The sensitivity of these animals to uranium exposure is unknown based on the existing scientific literature, and species-specific uranium presumptive effects levels were only available for two endangered fish species known to inhabit the segregation areas. Overall, the chemical toxicity data available for biological receptors of concern were limited, although chemical and radiation toxicity guidance values are available from several sources. However, caution should be used when directly applying these values to northern Arizona given the unique habitat and life history strategies of biological receptors in the segregation areas and the fact that some guidance values are based on models rather than empirical (laboratory or field) data. No chemical toxicity information based on empirical data is available for reptiles, birds, or wild mammals; therefore, the risks associated with uranium and other radionuclides are unknown for these biota.

Arizona↗

USGS2019

Synthesis of maternal transfer of mercury in birds: Implications for altered toxicity risk

Maternal transfer is the predominant route of methylmercury (MeHg) exposure to offspring. We reviewed and synthesized published and unpublished data on maternal transfer of MeHg in birds. Using paired samples of females’ blood (n=564) and their eggs (n=1814) from 26 bird species in 6 taxonomic orders, we conducted a meta-analysis to evaluate whether maternal transfer of MeHg to eggs differed among species and caused differential toxicity risk to embryos. Total mercury (THg) concentrations in eggs increased with maternal blood THg concentrations; however, the proportion of THg transferred from females to their eggs differed among bird taxa and with maternal THg exposure. Specifically, a smaller proportion of maternal THg was transferred to eggs with increasing female THg concentrations. Additionally, the proportion of THg that was transferred to eggs at the same maternal blood THg concentration differed among taxonomic orders, with waterfowl (Anseriformes) transferring up to 382% more THg into their eggs than songbirds (Passeriformes). We provide equations to predict THg concentrations in eggs using female blood THg concentrations, and vice versa, which may help translate toxicity benchmarks across tissues and life stages. Our results indicate that toxicity risk of MeHg can vary among bird taxa due to differences in the maternal transfer of MeHg to offspring.

Environmental Science & Technology↗

USGS2014

Carryover effects and climatic conditions influence the postfledging survival of greater sage-grouse

Prebreeding survival is an important life history component that affects both parental fitness and population persistence. In birds, prebreeding can be separated into pre- and postfledging periods; carryover effects from the prefledging period may influence postfledging survival. We investigated effects of body condition at fledging, and climatic variation, on postfledging survival of radio-marked greater sage-grouse ( Centrocercus urophasianus ) in the Great Basin Desert of the western United States. We hypothesized that body condition would influence postfledging survival as a carryover effect from the prefledging period, and we predicted that climatic variation may mediate this carryover effect or, alternatively, would act directly on survival during the postfledging period. Individual body condition had a strong positive effect on postfledging survival of juvenile females, suggesting carryover effects from the prefledging period. Females in the upper 25th percentile of body condition scores had a postfledging survival probability more than twice that (Φ = 0.51 ± 0.06 SE) of females in the bottom 25th percentile (Φ = 0.21 ± 0.05 SE). A similar effect could not be detected for males. We also found evidence for temperature and precipitation effects on monthly survival rates of both sexes. After controlling for site-level variation, postfledging survival was nearly twice as great following the coolest and wettest growing season (Φ = 0.77 ± 0.05 SE) compared with the hottest and driest growing season (Φ = 0.39 ± 0.05 SE). We found no relationships between individual body condition and temperature or precipitation, suggesting that carryover effects operated independently of background climatic variation. The temperature and precipitation effects we observed likely produced a direct effect on mortality risk during the postfledging period. Conservation actions that focus on improving prefledging habitat for sage-grouse may have indirect benefits to survival during postfledging, due to carryover effects between the two life phases.

Ecology and Evolution↗

USGS2008

Field Marks of a Celebration: Roger Tory Peterson's Centennial Birthday

A red letter day in my life was April 27, 1934, the day I first met Roger. A birding friend, Elisha Atkins, had invited Clinton Reynolds and me to dinner to meet a famous ornithologist. We would all be going on a field trip to Newburyport on the Massachusetts coast the next day. The dinner conversation revolved about a new field guide that Mr. Peterson had just completed and that would be available in a few days. I couldn?t wait to see it! I had been birding since 1930, keying out live birds with Chapman?s Handbook of Birds of Eastern North America (1912) and Hoffman?s Guide to the Birds of New England and Eastern New York (1904). Both books had extensive keys based on color, size, bill shape and season, and pictures of heads or feet of some species. Positive bird identification was a long and tedious process. The field trip the next day with Roger was memorable, not for finding any rare or unusual birds, but for learning how to identify birds to species at a single good glance. I recall asking Roger if he could find a ring-billed gull among a group of gulls resting on a roof beside the Merrimac River. He immediately said, ?No there aren?t any ring-bills there; they would be immediately apparent by their slimmer shape.? There was no need to check the foot color on each bird. While it is easy to say that Roger revolutionized field guides, I truly believe there are few people worldwide under the age of 90 who can really appreciate the difference between the old way of keying out birds and the instant recognition promoted by the Peterson system. Today we take for granted that amateurs can identify birds accurately. Monitoring bird populations by Breeding Bird Surveys, atlas studies, Breeding Bird Censuses, migration banding, and many other studies relies on it. None of these would be possible if we were still keying out live birds using books designed to identify dead birds in the hand.

Audubon Naturalist News↗

USGS2023

Investing in a nest egg: Intraspecific variation in the timing of egg laying across a latitudinal gradient

Avian reproductive strategies vary widely, and many studies of life-history variation have focused on the incubation and hatching stages of nesting. Birds make proximate decisions regarding reproductive investment during the laying stage, and these decisions likely constrain and tradeoff with other traits and subsequent behaviors. However, we know relatively little about egg-laying stage behaviors given the difficulty of locating and monitoring nest sites from the onset of laying. We used non-invasive continuous video recording to quantify variation in the egg-laying behaviors of burrowing owls ( Athene cunicularia) along a 1400-km latitudinal gradient in western North America. Burrowing owls laid eggs disproportionately in the morning hours, and that tendency was strongest among first eggs in a clutch. However, selection appeared to act more strongly on laying intervals (the time between laying of consecutive eggs) than on the diel time of laying, and laying intervals varied widely among and within clutches. Laying intervals declined seasonally and with increasing clutch size but increased with increasing burrow temperature and as a function of laying stage nest attentiveness, which may be a strategy to preserve egg viability. Laying interval was positively correlated with the duration of hatching intervals, suggesting that laying interval duration is one mechanism (along with timing of incubation onset) that generates variation in hatching asynchrony. Our results lend support to two general hypotheses to explain laying schedules; selection favors laying eggs in the morning, but other selective pressures may override that pattern. These conclusions indicate that allocation decisions during laying are an important part of avian life-history strategies which are subject to energetic constraints and tradeoffs with other traits.

Oecologia↗