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USGS2003

Establishing appropriate measures for monitoring aging in birds: comparing short and long lived species

In order to reveal patterns of reproductive aging in birds we focus on a short lived species, the Japanese quail and the American kestrel, which has a life span of medium length. Quail have been studied extensively in the laboratory as models for understanding avian endocrinology and behavior, and as a subject for toxicological research and testing. In the lab, Japanese quail show age-related deterioration in endocrine, behavioral, and sensory system responses; the American kestrel is relatively long lived and shows moderate evidence of senescence in the oldest birds. Using data collected from captive kestrels at the Patuxent Wildlife Research Center, a database was designed to document selected parameters over the life cycle of the kestrels. Life table data collated from many species indicate that longer lived species of birds show senescence in survival ability but this pattern has not been established for reproductive function. We suggest that useful comparisons among species can be made by identifying stages in reproductive life history, organized on a relative time scale. Preliminary data from quail and kestrels, admittedly only two species, do not yet indicate a pattern of greater reproductive senescence in longer-lived birds.

Experimental Gerontology↗

USGS2009

Inferences about landbird abundance from count data: recent advances and future directions

We summarize results of a November 2006 workshop dealing with recent research on the estimation of landbird abundance from count data. Our conceptual framework includes a decomposition of the probability of detecting a bird potentially exposed to sampling efforts into four separate probabilities. Primary inference methods are described and include distance sampling, multiple observers, time of detection, and repeated counts. The detection parameters estimated by these different approaches differ, leading to different interpretations of resulting estimates of density and abundance. Simultaneous use of combinations of these different inference approaches can not only lead to increased precision but also provides the ability to decompose components of the detection process. Recent efforts to test the efficacy of these different approaches using natural systems and a new bird radio test system provide sobering conclusions about the ability of observers to detect and localize birds in auditory surveys. Recent research is reported on efforts to deal with such potential sources of error as bird misclassification, measurement error, and density gradients. Methods for inference about spatial and temporal variation in avian abundance are outlined. Discussion topics include opinions about the need to estimate detection probability when drawing inference about avian abundance, methodological recommendations based on the current state of knowledge and suggestions for future research.

Book chapter↗

USGS2021

Domoic acid and saxitoxin in seabirds in the United States between 2007 and 2018

As harmful algal blooms (HABs) increase in magnitude and duration worldwide, they are becoming an expanding threat to marine wildlife. Over the past decade, blooms of algae that produce the neurotoxins domoic acid (DA) and saxitoxin (STX) and documented concurrent seabird mortality events have increased bicoastally in the United States. We conducted a retrospective analysis of HAB related mortality events in California, Washington, and Rhode Island between 2007 and 2018 involving 12 species of seabirds, to document the levels, ranges, and patterns of DA and STX in eight sample types (kidney, liver, stomach, intestinal, cloacal, cecal contents, bile, blood) collected from birds during these events. Samples ( n = 182 ) from 83 birds were examined for DA ( n = 135) or STX ( n = 17) or both toxins simultaneously ( n = 30), using ELISA or LCMS at the National Oceanographic and Atmospheric Administration, National Marine Fisheries Service (NOAA-NMFS) Wildlife Algal-toxin Research and Response Network (WARRN-West) or the University of California, Santa Cruz (UCSC). DA or STX was detected in seven of the sample types with STX below the minimum detection limit in blood for the three samples tested. DA was found in 70% and STX was found in 23% of all tested samples. The ranges of detectable levels of DA and STX in all samples were 0.65–681,190.00 ng g −1 and 2.00–20.95 ng g −1 , respectively. Cloacal contents from a Pacific loon ( Gavia pacifica ) collected in 2017 from Ventura County, California, had the highest maximum level of DA for all samples and species tested in this study. The highest level of STX for all samples and species was detected in the bile of a northern fulmar ( Fulmarus glacialis ) collected in 2018 from San Luis Obispo County, California. DA detections were consistently found in gastrointestinal samples, liver, bile, and kidney, whereas STX detections were most frequently seen in liver and bile samples. Co-occurring HAB toxins (DA and STX) were detected in white-winged scoters ( Melanitta deglandi ) in 2009, a Brandt's cormorant ( Phalacrocorax penicillatus ) in 2015, and a northern fulmar and common murre ( Uria aalge ) in 2018. This article provides DA and STX tissue concentrations and patterns in avian samples and shows the utility of various sample types for the detection of HAB toxins. Future research to understand the pharmacodynamics of these toxins in avian species and to establish lethal doses in various bird species would be beneficial.

Harmful Algae↗

USGS2015

Statistical guidelines for assessing marine avian hotspots and coldspots: A case study on wind energy development in the U.S. Atlantic Ocean

Estimating patterns of habitat use is challenging for marine avian species because seabirds tend to aggregate in large groups and it can be difficult to locate both individuals and groups in vast marine environments. We developed an approach to estimate the statistical power of discrete survey events to identify species-specific hotspots and coldspots of long-term seabird abundance in marine environments. We illustrate our approach using historical seabird data from survey transects in the U.S. Atlantic Ocean Outer Continental Shelf (OCS), an area that has been divided into “lease blocks” for proposed offshore wind energy development. For our power analysis, we examined whether discrete lease blocks within the region could be defined as hotspots (3 × mean abundance in the OCS) or coldspots (1/3 ×) for individual species within a given season. For each of 74 species/season combinations, we determined which of eight candidate statistical distributions (ranging in their degree of skewedness) best fit the count data. We then used the selected distribution and estimates of regional prevalence to calculate and map statistical power to detect hotspots and coldspots, and estimate the p -value from Monte Carlo significance tests that specific lease blocks are in fact hotspots or coldspots relative to regional average abundance. The power to detect species-specific hotspots was higher than that of coldspots for most species because species-specific prevalence was relatively low (mean: 0.111; SD: 0.110). The number of surveys required for adequate power (> 0.6) was large for most species (tens to hundreds) using this hotspot definition. Regulators may need to accept higher proportional effect sizes, combine species into groups, and/or broaden the spatial scale by combining lease blocks in order to determine optimal placement of wind farms. Our power analysis approach provides a general framework for both retrospective analyses and future avian survey design and is applicable to a broad range of research and conservation problems.

U.S. Atlantic Ocean Outer Continental Shelf↗

USGS2018

Preliminary evaluation of behavioral response of nesting waterbirds to small unmanned aircraft flight

Small unmanned aircraft systems present an emerging technology with the potential to survey colonial waterbird populations while reducing disturbance in comparison to traditional ground counts. Recent research with these systems has been performed on some colonially nesting avian species; however, none have focused on wading bird species. During 2015–2016, this study tested the behavioral response of a mixed-species rookery (Cattle Egret ( Bubulcus ibis ), Snowy Egret ( Egretta thula ), Glossy Ibis ( Plegadis falcinellus ) and a groundnesting colony of Common Terns ( Sterna hirundo )) in shrub habitat to small unmanned aircraft system flights at 12 m, 15 m, 30 m, and 50 m. Even at the lowest altitudes, the birds either showed no reaction or acclimated within 60 sec of the fly-over. Conversely, physically entering the colony to conduct ground surveys resulted in all Common Terns flushing from their nests beginning when the observer was 50 m away and required significantly more time in the colony overall: ~30–60 min vs. ~3–7 min with the small unmanned aircraft system. While this study focuses only on the behavioral response of nesting birds and not comparison of count estimates, these results provide preliminary evidence that small unmanned aircraft systems provide the potential to monitor colonial nesting bird populations while minimizing disturbance to the colony.

Waterbirds↗

USGS1994

A glossary for avian conservation biology

This glossary provides standard definitions for many of the terms used in avian conservation biology. We compiled these definitions to assist communication among researchers, managers, and others involved in the Neotropical Migratory Bird Conservation Program, also known as Partners in Flight. We used existing glossaries and recent literature to prepare this glossary. The cited sources were not necessarily the first ones to use the terms. Many definitions were taken verbatim from the cited source material. Others were modified slightly to clarify the meaning. Definitions that were modified to a greater extent are indicated as being adapted from the originals. Terms that have been used in more than one way by different authors are listed with numbered alternative definitions if the definitions differ substantially.

The Wilson Bulletin↗

USGS2021

An inventory and typology of permanent floodplain lakes in the Mississippi alluvial valley: A first step to conservation planning

The alluvial valley of the Mississippi River is an extensive area harboring hundreds of lakes created by fluvial dynamics. These floodplain lakes are scattered throughout the valley and carved over thousands of years by shifting river courses and other hydro-fluvial processes associated with contemporary and prehistoric rivers. These lakes have significant ecological importance as they support a large component of North American biodiversity. We used remote sensing to catalog lakes, to characterize morphology, and to construct a typology via cluster analysis. We identified over 1,300 permanent lakes totaling over 100,000 ha. The lakes were classified into 12 types according to lake size, shape, depth, connectivity, inundation frequency, and surrounding landcover. We anticipate that biotic characteristics differ among the 12 types, but large-scale systematic analyses of biotic assemblages of floodplain lakes in the region are mostly absent. Our typology can provide the framework essential for organizing research to define water dynamics, water quality, and ecological conditions such as forests, mussel, fish, and avian communities to construct conservation plans. The typology encourages a large-scale view of the properties of floodplain lakes in the alluvial valley. It is a functional tool that can be used to begin identifying conservation and research needs, adapt monitoring and management programs, customize environmental programs, and use conservation resources more effectively to achieve large-scale management objectives.

Mississippi River↗

USGS2010

Victims and vectors: highly pathogenic avian influenza H5N1 and the ecology of wild birds

The emergence of highly pathogenic avian influenza (HPAI) viruses has raised concerns about the role of wild birds in the spread and persistence of the disease. In 2005, an outbreak of the highly pathogenic subtype H5N1 killed more than 6,000 wild waterbirds at Qinghai Lake, China. Outbreaks have continued to periodically occur in wild birds at Qinghai Lake and elsewhere in Central China and Mongolia. This region has few poultry but is a major migration and breeding area for waterbirds in the Central Asian Flyway, although relatively little is known about migratory movements of different species and connectivity of their wetland habitats. The scientific debate has focused on the role of waterbirds in the epidemiology, maintenance and spread of HPAI H5N1: to what extent are they victims affected by the disease, or vectors that have a role in disease transmission? In this review, we summarise the current knowledge of wild bird involvement in the ecology of HPAI H5N1. Specifically, we present details on: (1) origin of HPAI H5N1; (2) waterbirds as LPAI reservoirs and evolution into HPAI; (3) the role of waterbirds in virus spread and persistence; (4) key biogeographic regions of outbreak; and (5) applying an ecological research perspective to studying AIVs in wild waterbirds and their ecosystems.

Avian Biology Research↗

USGS2018

Temperature-influenced energetics model for migrating waterfowl

Climate and weather affect avian migration by influencing when and where birds fly, the energy costs and risks of flight, and the ability to sense cues necessary for proper navigation. We review the literature of the physiology of avian migration and the influence of climate, specifically temperature, on avian migration dynamics. We use waterfowl as a model guild because of the ready availability of empirical physiological data and their enormous economic value, but our discussion and expectations are broadly generalizable to migratory birds in general. We detail potential consequences of an increasingly warm climate on avian migration, including the possibility of the cessation of migration by some populations and species. Our intent is to lay the groundwork for including temperature effects on energetic gains and losses of migratory birds with the expected consequences of increasing temperatures into a predictive modeling framework. To this end, we provide a simulation of migration progression exclusively focused on the influence of temperature on the physiological determinants of migration. This simulation produced comparable results to empirically derived and observed values for different migratory factors (e.g., body fat content, flight range, departure date). By merging knowledge from the arenas of avian physiology and migratory theory we have identified a clear need for research and have developed hypotheses for a path forward.

Ecological Modelling↗

USGS2022

Underwater hearing in sea ducks with applications for reducing gillnet bycatch through acoustic deterrence

As diving foragers, sea ducks are vulnerable to underwater anthropogenic activity, including ships, underwater construction, seismic surveys and gillnet fisheries. Bycatch in gillnets is a contributing source of mortality for sea ducks, killing hundreds of thousands of individuals annually. We researched underwater hearing in sea duck species to increase knowledge of underwater avian acoustic sensitivity and to assist with possible development of gillnet bycatch mitigation strategies that include auditory deterrent devices. We used both psychoacoustic and electrophysiological techniques to investigate underwater duck hearing in several species including the long-tailed duck ( Clangula hyemalis ), surf scoter ( Melanitta perspicillata ) and common eider ( Somateria mollissima ). Psychoacoustic results demonstrated that all species tested share a common range of maximum auditory sensitivity of 1.0–3.0 kHz, with the long-tailed ducks and common eiders at the high end of that range (2.96 kHz), and surf scoters at the low end (1.0 kHz). In addition, our electrophysiological results from 4 surf scoters and 2 long-tailed ducks, while only tested at 0.5, 1 and 2 kHz, generally agree with the audiogram shape from our psychoacoustic testing. The results from this study are applicable to the development of effective acoustic deterrent devices or pingers in the 2–3 kHz range to deter sea ducks from anthropogenic threats.

Journal of Experimental Biology↗

USGS2022

Avian predation on juvenile and adult Lost River and Shortnose Suckers: An updated multi-predator species evaluation

Previous research suggests that predation by piscivorous colonial waterbirds may negatively influence the survival of Lost River Suckers (LRS) Deltistes luxatus and Shortnose Suckers (SNS) Chasmistes brevirostris in the Upper Klamath Basin (UKB), USA. However, estimates of predation from past studies, which were based on suckers with PIT tags, represent minimum estimates of sucker mortality because analyses did not account for the proportion of tags that were consumed by birds and deposited beyond their breeding colony. To address this uncertainty, we fed PIT-tagged suckers to American white pelicans Pelecanus erythrorhynchos to estimate deposition probabilities. A hierarchical Bayesian model was then used to estimate predation rates (percentage of available tagged fish that were consumed) on juvenile suckers that were released as part of the Sucker Assisted Rearing Program (SARP) and on wild juvenile and adult LRS and SNS during 2009–2020. Pelican deposition probabilities were estimated to be 0.47 (95% credible interval = 0.36–0.60), indicating that for every 100 tags consumed, 47 tags on average were deposited on breeding colonies by birds. Deposition-corrected estimates of predation rates were approximately two times greater than those previously reported and ranged annually from 4.3% (95% credible interval = 2.9–6.7%) to 8.5% (6.3–12.7%) on SARP juvenile suckers, from 4.3% (0.9–13.2%) to 10.5% (3.8–24.5%) on wild juvenile suckers, and from 0.1% (<0.1–0.3%) to 7.2% (2.8–16.4%) on adult suckers, depending on species and location. Results suggest that predation by colonial waterbirds, although not the original cause of sucker declines, was a substantial source of sucker mortality in some years. Future studies should consider models that jointly estimate both predation and survival and models that include environmental factors that potentially influence sucker susceptibility to avian predators in the UKB.

California, Oregon↗

USGS2026

White light reduces flight height of birds in low-density migration corridor

Artificial light at night (ALAN) has been shown to influence the behavior of migratory birds, yet how different light spectra modulate these effects remains unclear. We conducted a field experiment for 31 nights at a remote site in northern Colorado, USA, using 2 800-W light-emitting diode (LED) floodlights with white, red, amber, and blue lighting treatments during fall migration in 2023 and 2024. Using a vertical-looking radar system, we quantified avian in-flight responses in migration traffic rate, flight height, and flight direction between paired control periods with no light and periods with lighting treatments. Although we found no effects of our treatments on migration traffic rate and weak effects on flight direction, we found that broad-spectrum white light with emissions across a 400–600 nm spectrum reduced mean avian flight height by an estimated 22.57 m (±10.53 SE) compared to dark periods, and this response was stronger than red and blue lighting treatments. Our ability to detect behavior changes from a point source in a rural, low-density migration system supports the notion that the effects of artificial light at night on bird behavior may be more pervasive than is often recognized, and that this varies by ALAN spectra. Beyond providing insight into avian orientation and in-flight behavior, our results have implications for future research, the conservation management of artificial light at night, and lighting designs that minimize effects on migratory birds.

Colorado↗

USGS2026

Bird migration and energetics simulations incorporating oil spill effects

Oil spills are well-known for causing acute mortality of birds, but sublethal and delayed impacts are less understood. Focusing on the mallard ( Anas platyrhynchos ), we used simulation modeling to explore how sublethal oiling may affect avian survival and breeding ground body condition. We used empirically informed migration and energetics simulations to model hypothetical spills occurring in northern Arkansas, USA occurring in either January to simulate thermoregulatory stress or March to simulate pre-migration effects. We modeled trace and lightly oiled female mallards (≤5% or 6 to 20% of feather area oiled, respectively), incorporating oiling-induced energetic effects on thermoregulation, flight, and energetic gain. We found that mortality was generally higher for simulated spills occurring in January versus March. In the simulations, mallards lost body mass due to oiling, but surviving individuals could partially recover body mass before arriving at the breeding grounds. Including oiling-induced energetic gain effects in simulations increased mortality as well as increased overall variability of simulation results. This modeling effort identified an important gap in knowledge regarding oiled bird energetics, specifically a need to better quantify oiling-induced energetic gain changes. Although the model is currently limited to a specific species and geographic area, it serves as a proof-of-concept for future research and modeling efforts aimed at understanding more broadly the impacts of oil spills on avian populations.

Prairie Pothole region↗

USGS2026

Phenological flexibility in nesting grassland birds over a half century informs timing of management

The timing of biological events for wildlife species reflects and influences ecological and evolutionary processes and helps guide management actions to meet conservation objectives. Climate change may cause phenological shifts across trophic levels and ecosystems, which can have broad ecological impacts. Shifts may also result in mistiming between management practices based on historical conditions and current biological events, reducing the effectiveness of natural resource management and conservation efforts. We analyzed nesting phenology patterns over recent decades across North America for 44 species of grassland-nesting birds, including waterfowl, upland gamebirds, shorebirds, and passerines, many of which have experienced precipitous population declines. Understanding nesting phenologies of these species is important to their conservation, as grassland management activities often conflict with reproductive periods leading to high nest and young mortality. We compiled a dataset of over 172,000 nest records spanning five decades and evaluated environmental factors influencing long-term shifts and short-term flexibility in nesting phenology. We found considerable variation in responses among species, with differences in both direction and magnitude of spatiotemporal effects on nest initiation dates. Notably, the community-level nesting season lengthened over time, but shifts exhibited spatial heterogeneity within this community of grassland birds. Nesting dates shifted asymmetrically across the breeding season with differential changes in early, peak, and late nesting dates whereas long-term changes were mediated by weather and geographic gradients. For example, nest initiation was earlier in warmer and wetter years. By applying a quantile-based analysis, we provide estimates of the full nesting period that could serve as a flexible tool for scheduling monitoring efforts and balancing management and human activities with periods of reduced nesting activity. Our collaborative effort illustrates the challenges and opportunities of collecting long-term data on avian communities necessary for answering questions across large regions but highlights the value of research coordination for bird conservation.

Global Change Biology↗

USGS2023

Disparate data streams together yield novel survival estimates of Alaska-breeding Whimbrels

Survival estimates are critical components of avian ecology. In well-intentioned efforts to maximize the utility of one's research, survival estimates often derive from data that were not originally collected for survival assessments, and such post hoc analyses may include unintentional biases. We estimated the survival of Whimbrels captured and marked at two breeding sites in Alaska using divergent data streams that in isolation were subject to methodological biases. Although both capture sites were chosen to study the migration ecology of Alaska-breeding Whimbrels, maximizing the conservation value of the data we collected was obviously desirable. We used multi-year telemetry information to infer survival from one site (Colville River) and mark-resight techniques to estimate survival from a second site (Kanuti River). At the Colville River, we could not feasibly include a control group of birds to assess potential survival effects of externally mounted transmitters, while at Kanuti River we were unable to accurately account for potential emigration events because we used resightings alone. We integrated these datasets in a Bayesian hierarchical framework, an approach that permitted insights across sites that moderated methodological biases within sites. Using telemetry enabled us to detect permanent emigration events from breeding sites in two of ten birds; results that informed estimates for birds without tracking devices. These datasets yielded point estimates of true survival of Whimbrels from Colville River equipped with solar-powered satellite transmitters that were higher (0.83) than true survival estimates of Whimbrels from Kanuti River marked with leg flags alone (0.74) or equipped with surgically implanted satellite transmitters (0.50), but the 95% credible intervals on these estimates overlapped across groups. For species like Whimbrels that are difficult and costly to study, combining information from disparate data streams allowed us to derive novel demographic estimates, an approach with clear application to other similar studies.

Alaska↗

USGS2007

Applying the scientific method when assessing the influence of migratory birds on the dispersal of H5N1

Background The role of wild birds in the dispersal of highly pathogenic avian influenza virus H5N1 continues to be the subject of considerable debate. However, some researchers functionally examining the same question are applying opposing null hypotheses when examining this issue. Discussion I describe the correct method for establishing a null hypothesis under the scientific method. I suggest that the correct null hypothesis is that migratory birds can disperse this virus during migration and encourage researchers to design studies to falsify this null. Finally, I provide several examples where statements made during this debate, while strictly true, are not generally informative or are speculative. Summary By adhering to the scientific method, definitive answers regarding the role of wild birds in the dispersal of highly pathogenic viruses will be reached more effectively.

Virology Journal↗

USGS2017

The pathogenesis of clade 2.3.4.4 H5 highly pathogenic avian influenza viruses in Ruddy Duck (Oxyura jamaicensis) and Lesser Scaup (Aythya affinis)

Waterfowl are the natural hosts of avian influenza virus (AIV) and disseminate the virus worldwide through migration. Historically, surveillance and research efforts for AIV in waterfowl have focused on dabbling ducks. The role of diving ducks in AIV ecology has not been well characterized. In this study, we examined the relative susceptibility and pathogenicity of clade 2.3.4.4 H5 highly pathogenic AIV (HPAIV) in two species of diving ducks. Juvenile and adult Ruddy Duck ( Oxyura jamaicensis ) and juvenile Lesser Scaup ( Aythya affinis ) were intranasally inoculated with A/Northern Pintail/WA/40964/2014 H5N2 HPAIV. Additional groups of juvenile Lesser Scaups were inoculated with A/Gyrfalcon/WA/41088/2014 H5N8 HPAIV. The approximate 50% bird infectious doses (BID 50 ) of the H5N2 isolate for adult Ruddy Ducks was <10 2 50% egg infectious doses (EID 50 ) and for the juvenile Lesser Scaups it was <10 4 EID 50 . There were insufficient juvenile Ruddy Ducks to calculate the BID 50 . The BID 50 for the juvenile Lesser Scaups inoculated with the H5N8 isolate was 10 3 EID 50 . Clinical disease was not observed in any group; however, mortality occurred in the juvenile Ruddy Ducks inoculated with the H5N2 virus (three of five ducks), and staining for AIV antigen was observed in numerous tissues from these ducks. One adult Ruddy Duck also died and although it was infected with AIV (the duck was positive for virus shedding and AIV antigen was detected in tissues), it was also infected with coccidiosis. The proportion of ducks shedding virus was related to the dose administered, but the titers were similar among dose groups. The group with the fewest ducks shedding virus was the adult Ruddy Ducks. There was a trend for the Lesser Scaups to shed higher titers of virus than the Ruddy Ducks. No virus shedding was detected after 7 d postinoculation in any group. Similar to dabbling ducks, Lesser Scaups and Ruddy Ducks are susceptible to infection with this H5 HPAIV lineage, although they excrete lower titers of virus.

Journal of Wildlife Diseases↗

USGS2015

The effects of heterospecifics and climatic conditions on incubation behavior within a mixed-species colony

Parental incubation behavior largely influences nest survival, a critical demographic process in avian population dynamics, and behaviors vary across species with different life history breeding strategies. Although research has identified nest survival advantages of mixing colonies, behavioral mechanisms that might explain these effects is largely lacking. We examined parental incubation behavior using video-monitoring techniques on Alcatraz Island, California, of black-crowned night-heron Nycticorax nycticorax (hereinafter, night-heron) in a mixed-species colony with California gulls Larus californicus and western gulls L. occidentalis . We first quantified general nesting behaviors (i.e. incubation constancy, and nest attendance), and a suite of specific nesting behaviors (i.e. inactivity, vigilance, preening, and nest maintenance) with respect to six different daily time periods. We employed linear mixed effects models to investigate environmental and temporal factors as sources of variation in incubation constancy and nest attendance using 211 nest days across three nesting seasons (2010–2012). We found incubation constancy (percent of time on the eggs) and nest attendance (percent of time at the nest) were lower for nests that were located < 3 m from one or more gull nest, which indirectly supports the predator protection hypothesis, whereby heterospecifics provide protection allowing more time for foraging and other self-maintenance activities. To our knowledge, this is the first empirical evidence of the influence of one nesting species on the incubation behavior of another. We also identified distinct differences between incubation constancy and nest attentiveness, indicating that these biparental incubating species do not share similar energetic constraints as those that are observed for uniparental species. Additionally, we found that variation in incubation behavior was a function of temperature and precipitation, where the strength of these effects was dependent on the time of day. Overall, these findings strengthen our understanding of incubation behavior and nest ecology of a colonial-nesting species.

California↗