Timothy D. Bowman’s research while affiliated with U.S. Fish and Wildlife Service and other places

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Publications (14)


(a) Multispecies annual‐cycle network for North American black scoter (Melanitta americana), surf scoter (Melanitta perspicillata), and white‐winged scoter (Melanitta deglandi) showing seasonal use (blue = winter, green = spring migration, red = breeding, orange = fall migration), 1998–2019. Node sizes correspond to relative intensity of use, with the node size key (nested circles) depicting numbers of centroids for small‐, medium‐, and large‐diameter nodes. Inset maps depict areas of overlap between individuals from western (dark purple) and eastern (dark pink) capture sites for (b) white‐winged scoter and (c) surf scoter. Eastern and western black scoter populations did not overlap.
Multispecies values for (a) indegree centrality, (b) outdegree centrality, (c) betweenness centrality, and (d) connectivity for North American scoters, 1998–2019. For complete descriptions of the four network centrality metrics, see Appendix S1: Figure S1.
Network modularity for North American scoters tracked between 1998 and 2019 for (a) all three scoter species combined, (b) surf scoters only, (c) black scoters only, and (d) white‐winged scoters only. Colors indicate groups of nodes (ie modules) that are more interconnected with one another than with the overall network. Modularity values (Girvan‐Newman scores) ranged from 0 (random) to 1 (complete interconnection within modules and no connections between modules); we considered values >0.5 to represent high modularity. Dashed lines in (a) show boundaries of traditional waterfowl flyways (Pacific, Central, Mississippi, and Atlantic) in the US and Canada.
Evaluating conservation units using network analysis: a sea duck case study
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January 2024

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228 Reads

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4 Citations

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Scott G Gilliland

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[...]

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Conserving migratory wildlife requires understanding how groups of individuals interact across seasons and landscapes. Telemetry reveals individual movements at large spatiotemporal scales; however, using movement data to define conservation units requires scaling up from individual movements to species‐ and community‐level patterns. We developed a framework to define flyways and identify important sites from telemetry data and applied it to long‐term, range‐wide tracking data from three species (640 individuals) of sea ducks: namely, North American scoters (Melanitta spp). Our network of 88 nodes included both multispecies hotspots and areas uniquely important to individual species. We found limited spatial overlap between scoters wintering on the Atlantic and Pacific coasts of North America, with differing connectivity patterns between coasts. Finally, we identified four multispecies conservation units that did not correspond to traditional management flyways. From this approach, we show how individual movements can be used to quantify range‐wide connectivity of migratory species and reveal gaps in conservation strategies.

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Continental distribution of North American black scoter (BLSC), surf scoter (SUSC), and white‐winged scoter (WWSC) estimated nesting locations determined from PTT‐tagged birds marked on Atlantic or Pacific wintering, staging, and molt sites between 1999 and 2020. Background colours indicate Level I North American Ecoregions (included for illustration). Level 1 ecoregion shapefile from the US Environmental Protection Agency (USEPA 2010).
Estimated nesting location arrival dates of PTT‐marked scoters varied by species (BLSC = black scoter, SUSC = surf scoter, WWSC = white‐winged scoter) and wintering origin (Atlantic = green, Pacific = purple; a), age at breeding (b), snow‐free date (c), and sex (d). In panels (a, b and d), boxes show model‐predicted means ± SE, with vertical lines indicating 85% confidence intervals. In panel c, line indicates model‐predicted means ± an 85% confidence interval. In all panels, circles indicate raw values for individual birds. See Table 1–2 for sample sizes.
Map of model‐predicted nesting location arrival dates of black, surf, and white‐winged scoters. Predictions were made across a polygon covering a 100 km buffer around the Taiga and Hudson Plains ecoregions, extended to include PTT‐estimated scoter nesting locations along the northwest coast of Alaska. Predictions were generated using the 11‐year snow‐free date average from 2010–2020 within each cell of a hexagonal grid with 24 km apothems (i.e. distance from hexagon centre to side). Map colours indicate arrival date, with more blue colours indicating earlier arrivals, and more yellow colours indicating later arrivals. Note that maps do not depict predicted breeding ranges, as the map includes areas outside of the presumed breeding range of certain species (e.g. black scoters do not breed across the entire continent; Bordage and Savard 2020), and birds do not nest at every location within the area where predictions were made (e.g. on mountains and large lakes, along the St. Lawrence River shoreline).
Minimum length of stay of male PTT‐marked scoters at estimated nesting locations varied by snow‐free date. Line indicates model‐based predictions ± an 85% confidence interval. Circles indicate raw values for individual birds. See Table 1–2 for sample sizes.
Departure dates of male PTT‐marked scoters from estimated nesting locations varied by species (BLSC = black scoter, SUSC = surf scoter, WWSC = white‐winged scoter) and wintering origin (Atlantic = green, Pacific = purple; a) and age at breeding (b). Boxes show model‐predicted means ± SE, with vertical lines indicating 95% confidence intervals. See Table 1–2 for sample sizes.
Evaluation of breeding distribution and chronology of North American scoters

October 2023

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190 Reads

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3 Citations

North America's scoter species are poorly monitored relative to other waterfowl. Black Melanitta americana, surf M. perspicillata, and white‐winged M. deglandi scoter abundance and trend estimates are thus uncertain in many parts of these species' ranges. The most extensive source of waterfowl abundance and distribution data in North America is the Waterfowl breeding population and habitat survey (WBPHS). Although the WBPHS effectively monitors most species, both its timing and geographic coverage may preclude accurate scoter monitoring. Therefore, our goal was to better define when and where scoters breed to help interpret survey results and optimize potential supplemental survey efforts for scoters. We integrated satellite telemetry tracking data from scoters marked at multiple molting, staging, breeding, and wintering areas along the Atlantic and Pacific coasts to quantify continent‐wide breeding chronology and distribution. We also examined possible drivers of variation in timing of arrival, length of stay, and departure at nesting locations. We documented a northwest to southeast distribution of estimated breeding sites across Alaska and Canada. On average, scoters arrived at nest sites on 1 June. Surf scoters and Pacific black scoters arrived earliest and departed earliest. Pacific‐wintering black and white‐winged scoters began breeding earlier than Atlantic‐wintering birds. Additionally, birds arrived at nesting locations earlier in years with earlier snowmelt, and later snowmelt reduced lengths of stay for males. Breeding chronology also varied by age group, with adults arriving earlier than subadults. Our study is the first to comprehensively describe spatial variation in timing of breeding of both Atlantic and Pacific populations of all three scoter species across North America. Our results increase our understanding of how current surveys enumerate scoters and will inform possible supplemental efforts to improve continental monitoring of scoter populations.


Model selection table for candidate base and final models explaining variation in fall migration parameters. Included are number of estimated parameters (K), correct Akaike's information criterion (AIC c ), AIC c , model probabilities (Akaike weight, w i ), log-likelihood (LL), and Nakagawa's conditional and marginal R 2 . All models include a random effect of bird identity. Candidate models with ΔAIC c < 2 and null models are shown. For final models, base models are shown regardless of ΔAIC c value.
Pacific Barrow’s Goldeneye refine migratory phenology in response to overwintering temperatures and annual snow melt

May 2023

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78 Reads

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3 Citations

Ornithology

Timing of seasonal bird migrations is broadly determined by internal biological clocks, which are synchronized by photoperiod, but individuals often refine their migratory timing decisions in response to external factors. Using 11 years of satellite telemetry data, we show that Pacific Barrow’s Goldeneye (Bucephala islandica), at higher latitudes, initiated spring and molt migrations later and fall migration earlier than individuals at lower latitudes. We further show that individuals refined migratory timing in response to interannual variation in environmental conditions. Individual Barrow’s Goldeneye initiated spring migration earlier in years with warmer springs at their overwintering locations and concluded spring migration earlier in years with earlier annual snow melt on their breeding grounds. Because individuals respond to conditions both where they initiate and where they conclude spring migration, our results suggest that Barrow’s Goldeneye update their migratory decisions en route. For all three migrations in their annual cycle, birds delayed initiating migration if they had been captured and tagged prior to that migration. Birds that initiated migration late for their latitude were less likely to include a stopover and completed that migration faster, partially compensating for delayed departures. Our results are consistent with the hypothesis that Barrow’s Goldeneye use a combination of endogenous cues and environmental cues in migratory decision making. Sensitivity to environmental cues suggests that Barrow’s Goldeneye may have behavioural plasticity that is adaptive when faced with ongoing climate change.


Fig. 3. Wintering latitude and sex effects on the date of arrival, departure, and distance traveled by adult Barrow's Goldeneye to the subsequent stage for wintering (1,4,7), breeding (2,5,8), and molting stages (3,6,9).
Migratory connectivity and variation in migration phenology within the Pacific population of Barrow's Goldeneye (Bucephala islandica)

August 2022

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63 Reads

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5 Citations

Avian Conservation and Ecology

Effective management of migratory animals requires an understanding of individual movement patterns throughout the annual cycle. We used satellite transmitters to track adult Barrow’s Goldeneye (Bucephala islandica) captured at five wintering sites, one breeding site, and one molting site from across the species’ geographic range in western North America. The data were analyzed to assess the strength of migratory connectivity and determine latitudinal and sex effects on migration phenology. At the range-wide scale, migratory connectivity was high; cluster analyses showed subpopulations aggregated at each stage of the annual cycle. Barrow’s Goldeneye from all wintering latitudes traveled north and east to breed. Compared to females, males traveled farther north to molt and consequently traveled longer distances when returning to their wintering areas. Wintering latitude had little effect on migration distance but a large effect on the phenology of migration. Individuals that wintered at northern latitudes arrived on their wintering areas earlier and departed later than individuals that wintered farther south. Individuals that wintered at northern latitudes also arrived on and left their inland breeding areas later but left their molting sites earlier than individuals that wintered farther south. Sex also influenced the phenology of migration at all sites: males left their mates during incubation and consequently spent less time on breeding areas and more time on their molting and fall staging areas. The high level of migratory connectivity observed in this study suggests that the subpopulation of Barrow’s Goldeneye in south-central Alaska is demographically independent from subpopulations in southeast Alaska and British Columbia, and could be managed separately.



Annual‐Cycle Movements and Phenology of Black Scoters in Eastern North America

September 2021

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94 Reads

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4 Citations

Journal of Wildlife Management

Sea ducks exhibit complex movement patterns throughout their annual cycle; most species use distinct molting and staging sites during migration and disjunct breeding and wintering sites. Although research on black scoters (Melanitta americana) has investigated movements and habitat selection during winter, little is known about their annual‐cycle movements. We used satellite telemetry to identify individual variation in migratory routes and breeding areas for black scoters wintering along the Atlantic Coast, to assess migratory connectivity among wintering, staging, breeding, and molt sites, and to examine effects of breeding site attendance on movement patterns and phenology. Black scoters occupied wintering areas from Canadian Maritime provinces to the southeastern United States. Males used an average of 2.5 distinct winter areas compared to 1.1 areas for females, and within‐winter movements averaged 1,256 km/individual. Individuals used an average of 2.1 staging sites during the 45‐day pre‐breeding migration period, and almost all were detected in the Gulf of St. Lawrence. Males spent less time at breeding sites and departed them earlier than females. During post‐breeding migration, females took approximately 25 fewer days than males to migrate from breeding sites to molt and staging sites, and then wintering areas. Most individuals used molt sites in James and Hudson bays before migrating directly to coastal wintering sites, which took approximately 11 days and covered 1,524 km. Males tended to arrive at wintering areas 10 days earlier than females. Individuals wintering near one another did not breed closer together than expected by chance, suggesting weak spatial structuring of the Atlantic population. Females exhibited greater fidelity (4.5 km) to previously used breeding sites compared to males (60 km). A substantial number of birds bred west of Hudson Bay in the Barrenlands, suggesting this area is used more widely than believed previously. Hudson and James bays provided key habitat for black scoters that winter along the Atlantic Coast, with most individuals residing for >30% of their annual cycle in these bays. Relative to other species of sea duck along the Atlantic Coast, the Atlantic population of black scoter is more dispersed and mobile during winter but is more concentrated during migration. These results could have implications for future survey efforts designed to assess population trends of black scoters. © 2021 The Wildlife Society. We used satellite telemetry to assess annual‐cycle movements of a large sample of black scoters, a sea duck that breeds in remote boreal areas and winters on marine coasts. Although we found a high degree of similarity in migration routes and timing and high fidelity to breeding sites, individuals showed low fidelity to wintering and migratory staging sites and used multiple sites among and within years. Our findings highlight the importance of migratory staging and molt areas in the Gulf of St. Lawrence and Hudson and James bays and the difficulty of accurately censusing black scoters during non‐breeding.


Reliability of External Characteristics to Age Barrow's Goldeneye

September 2020

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26 Reads

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3 Citations

Accurate assignment of age class is critical for understanding most demographic processes. For waterfowl, most techniques for determining age class require birds in hand, reducing utility for quickly and efficiently sampling a large portion of the population. As an alternative, we sought to establish an observation‐based methodology, achievable in the field with standard optics, for determining age class of Barrow's goldeneyes (Bucephala islandica). We photographed heads, wings, and bellies of 232 Barrow's goldeneyes captured during late winter (February–April) of 2007–2015 along the north Pacific Coast. From these photographs, we focused on 5 external characteristics for both males and females, with binary states that putatively corresponded to 2 age classes—first‐year birds (<1 yr) and adults (>1 yr). For males, all 5 external traits (belly color, head color, eye color, facial crescent, median secondary coverts color) had binary states that were reliably distinguishable by observers. Moreover, all 5 external traits were highly predictive of age class (≥96% concordance between external vs. bursal‐derived ages), and novice observers, after receiving training, were able to accurately age 96% of first‐year and 99% of adult males. In contrast, patterns were weaker for females; putative external characteristics of female age class (belly color, bill radiance, bill blackness, eye color, median secondary coverts color) had 77–91% concordance with bursal‐derived age, compared with 96–100% for males, and observers misidentified age classes of 15% of females, compared with only 2% of males. Overall, age classes of male Barrow's goldeneyes were accurately and reliably distinguishable during winter based on several external characteristics, whereas those of females were not. Our technique may be used to estimate age composition of male Barrow's goldeneyes during winter, providing a useful metric for monitoring annual changes in adult‐to‐juvenile ratios and other important demographic parameters. © 2020 The Wildlife Society. Most techniques for assigning age class of waterfowl require birds in the hand to identify detailed or concealed traits, largely limiting their utility to species that are commonly captured or harvested. We developed guidelines to accurately estimate age class of wintering Barrow's goldeneyes via observational surveys, finding that ages of male goldeneyes were reliably distinguishable based on several external characteristics whereas ages of females were not.


Annual Winter Site Fidelity of Barrow's Goldeneyes in the Pacific

October 2019

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64 Reads

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8 Citations

Journal of Wildlife Management

Coastal regions on the Pacific north coast of North America provide important wintering habitat for many species of sea ducks. Although winter range and habitat preferences are well described for most species, fidelity to coastal wintering sites is generally undocumented. Fidelity is an important factor necessary for understanding interactions with coastal developments and activities and corresponding management strategies. We used data from Barrow's goldeneyes (Bucephala islandica), a sea duck that winters predominantly in nearshore habitats along the Pacific north coast, to investigate inter‐annual fidelity to, and intra‐annual fidelity within, coastal wintering sites. Between 2006 and 2015, we marked goldeneyes on breeding, molting, and wintering sites with satellite transmitters. We retained 4,931 locations in coastal habitats from 221 goldeneyes across 4 coastal regions for our analyses. These birds demonstrated high inter‐annual fidelity to coastal wintering sites; 75% of selected wintering sites were within 29 km of sites used the previous winter. Inter‐annual fidelity to wintering sites was similar between sex and age classes but differed by coastal region. Goldeneyes from southcentral Alaska, USA, expressed greater inter‐annual fidelity relative to birds from northern or southern British Columbia, Canada, and southeast Alaska. Goldeneyes also expressed high intra‐annual fidelity within wintering sites, with 75% of individuals averaging within‐season movements of ≤9 km. Intra‐annual fidelity was lesser for female than male goldeneyes but did not differ between hatch‐year and after‐hatch‐year birds. We found regional variation in intra‐annual fidelity, with goldeneyes from southcentral Alaska expressing greater intra‐annual fidelity compared to birds from other regions. High inter‐ and intra‐annual winter site fidelity by Barrow's goldeneyes suggests that, at a population level, habitat use is predictable and can be used to inform risk assessment or to evaluate factors affecting habitat choice. Also, low dispersal among wintering sites suggests that recovery from population perturbations, whether caused by natural or anthropogenic events, will be protracted. © 2019 The Wildlife Society. High inter‐ and intra‐annual site fidelity by Barrow's goldeneyes along the Pacific north coast suggests that, at a population level, habitat use is predictable and can be used for assessing risks or to evaluate factors affecting habitat choice. Low dispersal of goldeneyes among wintering sites suggests that recovery from population perturbations, whether caused by natural or anthropogenic events, will be protracted.


Figure 1. Intranasal administration of midazolam hydrochloride (5 mg) in a surf scoter (M. perspicillata) shortly after its capture. A 22-Ga soft catheter was gently inserted into the nasal cavity while the head of the bird was held vertically (beak up), and half of the drug was instilled into each nare. The vertical position of the head was held for 2 min to allow for absorption of the drug.
Figure 2. Graphical representation of changes in mortality at 14 and 30 days after release in surf scoters (M. perspicillata) implanted with intracoelomic transmitters in two different treatment groups: 5 mg of midazolam hydrochloride via intranasal route (midazolam group) or an equivalent volume of saline (control group) upon capture. At 30 days after release, mortality was significantly different (P ¼ 0.004) as indicated with the horizontal bar (*). Error bars indicate standard error of the mean.
Intranasal administration of midazolam hydrochloride improves survival in female surf scoters (melanitta perspicillata) surgically implanted with intracoelomic transmitters

March 2019

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150 Reads

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10 Citations

Journal of Zoo and Wildlife Medicine

The intracoelomic implantation of satellite transmitters is associated with lower survival in surf scoters (Melanitta perspicillata) compared with other species of diving ducks, potentially due to physiologic alterations following physical exertion and stress caused by handling and confinement. The effect of intranasal administration of midazolam hydrochloride on survival of surf scoters surgically implanted with intracelomic transmitters was evaluated. Shortly after their capture in Forestville (QC, Canada) in the fall of 2013, 26 randomly selected adult female surf scoters were administered midazolam hydrochloride (4.6-5.9 mg/kg) intranasally. The same volume of saline (1 mL) was given to another 26 adult female surf scoters as a control group. All birds were surgically implanted with an intracoelomic transmitter equipped with a percutaneous antenna. Transmitters were programmed to transmit 2 hr each day for 30 days after implantation, and mortality was estimated for each group using the telemetry data. The association between the administration of midazolam and survival was assessed while controlling for other factors such as body mass, transmitter-mass-to-body-mass ratio, hematocrit, total solids, and duration of surgery, anesthesia, and confinement. The odds of presumed death in the saline group were 5.3 times higher than in the midazolam group (95% confidence interval: 1.7, 19.0; P = 0.004). The presumed mortality at 30 days for the midazolam group (23%) was lower than for the saline group (61%). No other variable was significantly associated with survival. These results suggest that sedation with midazolam shortly after capture increased the postsurgical survival of female surf scoters surgically implanted with intracoelomic transmitters.


Visualizing Populations of North American Sea Ducks: Maps to Guide Research and Management Planning

January 2019

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510 Reads

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2 Citations


Citations (12)


... animal migration, area-based conservation, bird ringing, movement ecology, network analysis, protected areas, satellite tracking, shorebirds Migratory site networks based on modelled movements have been used to identify important sites for waterfowl and assess their spatial coverage by protected areas (Deboelpaep et al., 2022;Xu et al., 2020). Yet, it remains unclear whether network approaches based on empirical movement data are feasible at the flyway scale (but see Lamb et al., 2024), which may be an important step for achieving formal incorporation of connectivity metrics into important site assessments for migratory species (Davidson et al., 1999). ...

Reference:

Site‐level connectivity identified from multiple sources of movement data to inform conservation of a migratory bird
Evaluating conservation units using network analysis: a sea duck case study

... Table 1. Results of generalized linear mixed models fitted to aerial survey data of Common Scoter in the Danish Aalborg Bugt and German North Sea to estimate the relationships between abundance and water depth (m), number of flocks and number of occupied grid squares and month of survey (see Fig. 3 and Supporting Information Table S3 migration, while most females and males continue to breeding areas (Lamb et al. 2021, Bianchini et al. 2023. ...

Evaluation of breeding distribution and chronology of North American scoters

... covariate (Kemp et al., 2023;Morin et al., 2020). By condensing these four covariates into a single group, we reduced the number of possible candidate models and allowed for a focused assessment of the relative importance of lidar-derived structure covariates (Morin et al., 2020). ...

Pacific Barrow’s Goldeneye refine migratory phenology in response to overwintering temperatures and annual snow melt

Ornithology

... Burrard Inlet, which is part of the Salish Sea is recognized as one of the world's most important bird areas supporting globally vital bird populations (Bowman et al., 2022). It is especially important to sea ducks of the Pacific Flyway during wintering, staging, spring migration, and molting. ...

Atlas of Sea Duck Key Habitat Sites in North America
  • Citing Technical Report
  • March 2022

... Our calculations of migratory connectivity without subadults resulted in higher values than those including all age groups. Numerous contemporary studies on migratory connectivity have concentrated on adults only (Forstner et al., 2022;Hill & Renfrew, 2019;Piironen et al., 2022) or neglected the age structure within populations (Huysman et al., 2022), potentially leading to biased estimations. Therefore, incorporating diverse age groups into migratory connectivity calculations is highly recommended for improving result accuracy. ...

Migratory connectivity and variation in migration phenology within the Pacific population of Barrow's Goldeneye (Bucephala islandica)

Avian Conservation and Ecology

... Table 1. Results of generalized linear mixed models fitted to aerial survey data of Common Scoter in the Danish Aalborg Bugt and German North Sea to estimate the relationships between abundance and water depth (m), number of flocks and number of occupied grid squares and month of survey (see Fig. 3 and Supporting Information Table S3 migration, while most females and males continue to breeding areas (Lamb et al. 2021, Bianchini et al. 2023. ...

Annual‐Cycle Movements and Phenology of Black Scoters in Eastern North America
  • Citing Article
  • September 2021

Journal of Wildlife Management

... The HY birds were almost fully grown but still flightless at ~ 6 weeks of age when captured and tagged. We recorded mass and wing chord of each bird and estimated sex and age class based on plumage [35], cloacal characteristics, bursal depth [36], and by comparing masses and wing sizes within each family cohort, under the assumption that males are larger at fledging than females [30]. ...

Reliability of External Characteristics to Age Barrow's Goldeneye
  • Citing Article
  • September 2020

... Interpretations of sea duck species genetic data benefit from the knowledge of species breeding ecology (Ely & Scribner 1994, juxtaposition of breeding, moulting and wintering habitats (Robertson & Cooke 1999, Robert et al. 2008, evidence of sex-biased gene flow (Scribner et al. 2001), site fidelity and migratory connectivity (Forstner et al. 2022), and the detection of historical refugia and subsequent post-glacial colonization (Ploeger 1968, Sonsthagen et al. 2011). Among sea duck species there is high variability in some of these traits, which would lead to different predictions about the occurrence and spatial scale of population differentiation (Talbot et al. 2015, Pearce et al. 2019. Background data are particularly relevant to forecast future changes in distribution and demography in the current era of climate variability and change (McKeon et al. 2016, CAFF 2017. ...

Visualizing Populations of North American Sea Ducks: Maps to Guide Research and Management Planning

... Because of strong site fidelity and differences in winter body condition, redheads from different parts of the Gulf of Mexico presumably represent different subpopulations and begin spring migration with varying nutrient reserves. Pintails in Texas coastal areas have also exhibited strong winter site fidelity (Hestbeck 1993), as have Barrow's goldeneyes Bucephala islandica along the Pacific coast (Willie et al. 2020). Mallards from four separate winter areas in eastern Colorado also showed different condition values, suggesting strong site fidelity (Ringelman and Szymczak 1985). ...

Annual Winter Site Fidelity of Barrow's Goldeneyes in the Pacific
  • Citing Article
  • October 2019

Journal of Wildlife Management

... Midazolam is a benzodiazepine used for chemical restraint of birds. It does not exhibit marked cardiovascular effects and provides satisfactory sedation with good muscle relaxation (Net et al., 2019). Usually, midazolam is used in combination with ketamine to provide greater sedation. ...

Intranasal administration of midazolam hydrochloride improves survival in female surf scoters (melanitta perspicillata) surgically implanted with intracoelomic transmitters

Journal of Zoo and Wildlife Medicine