Density-dependence produces spurious relationships among demographic parameters in a harvested species.

Harvest of wild organisms is an important component of human culture, economy, and recreation, but can also put species at risk of extinction. Decisions that guide successful management actions therefore rely on the ability of researchers to link changes in demographic processes to the anthropogenic actions or environmental changes that underlie variation in demographic parameters. Ecologists often use population models or maximum sustained yield curves to estimate the impacts of harvest on wildlife and fish populations. Applications of these models usually focus exclusively on the impact of harvest and often fail to consider adequately other potential, often collinear, mechanistic drivers of the observed relationships between harvest and demographic rates. In this study, we used an integrated population model and long-term data (1973-2016) to examine the relationships among hunting and natural mortality, the number of hunters, habitat conditions, and population size of blue-winged teal Spatula discors, an abundant North American dabbling duck with a relatively fast-paced life history strategy. Over the last two and a half decades of the study, teal abundance tripled, hunting mortality probability increased slightly ( < 0.02 ), and natural mortality probability increased substantially ( > 0.1 ) at greater population densities. We demonstrate strong density-dependent effects on natural mortality and fecundity as population density increased, indicative of compensatory harvest mortality and compensatory natality. Critically, an analysis that only assessed the relationship between survival and hunting mortality would spuriously indicate depensatory mortality due to multicollinearity between abundance, natural mortality and hunting mortality. Our findings demonstrate that models that only consider the direct effect of hunting on survival or natural mortality can fail to accurately assess the mechanistic impact of hunting on population dynamics due to multicollinearity among demographic drivers. This multicollinearity limits inference and may have strong impacts on applied management actions globally.

Informing wetland management with waterfowl movement and sanctuary use responses to human-induced disturbance.

Long-term environmental management to prevent waterfowl population declines is informed by ecology, movement behavior and habitat use patterns. Extrinsic factors, such as human-induced disturbance, can cause behavioral changes which may influence movement and resource needs, driving variation that affects management efficacy. To better understand the relationship between human-based disturbance and animal movement and habitat use, and their potential effects on management, we GPS tracked 15 dabbling ducks in California over ~4-weeks before, during and after the start of a recreational hunting season in October/November 2018. We recorded locations at 2-min intervals across three separate 24-h tracking phases: Phase 1) two weeks before the start of the hunting season (control (undisturbed) movement); Phase 2) the hunting season opening weekend; and Phase 3) a hunting weekend two weeks after opening weekend. We used GLMM models to analyze variation in movement and habitat use under hunting pressure compared with 'normal' observed patterns prior to commencement of hunting. We also compared responses to differing levels of disturbance related to the time of day (high - shooting/~daytime); moderate - non-lethal (~crepuscular); and low - night). During opening weekend flight (% time and distance) more than doubled during moderate and low disturbance and increased by ~50% during high disturbance compared with the pre-season weekend. Sanctuary use tripled during moderate and low disturbance and increased ~50% during high disturbance. Two weeks later flight decreased in all disturbance levels but was only less than the pre-season levels during high disturbance. In contrast, sanctuary use only decreased at night, although not to pre-season levels, while daytime doubled from ~45% to >80%. Birds adjust rapidly to disturbance and our results have implications for energetics models that estimate population food requirements. Management would benefit from reassessing the juxtaposition of essential sanctuary and feeding habitats to optimize wetland management for waterfowl.

Nocturnal incubation recess and flushing behavior by duck hens.

Incubating birds must balance the needs of their developing embryos with their own physiological needs, and many birds accomplish this by taking periodic breaks from incubation. Mallard (Anas platyrhynchos) and gadwall (Mareca strepera) hens typically take incubation recesses in the early morning and late afternoon, but recesses can also take place at night. We examined nocturnal incubation recess behavior for mallard and gadwall hens nesting in Suisun Marsh, California, USA, using iButton temperature dataloggers and continuous video monitoring at nests. Fourteen percent of all detected incubation recesses (N = 13,708) were nocturnal and took place on 20% of nest-days (N = 8,668). Video monitoring showed that hens covered their eggs with down feathers when they initiated a nocturnal recess themselves as they would a diurnal recess, but they left the eggs uncovered in 94% of the nocturnal recesses in which predators appeared at nests. Thus, determining whether or not eggs were left uncovered during a recess can provide strong indication whether the recess was initiated by the hen (eggs covered) or a predator (eggs uncovered). Because nest temperature decreased more rapidly when eggs were left uncovered versus covered, we were able to characterize eggs during nocturnal incubation recesses as covered or uncovered using nest temperature data. Overall, we predicted that 75% of nocturnal recesses were hen-initiated recesses (eggs covered) whereas 25% of nocturnal recesses were predator-initiated recesses (eggs uncovered). Of the predator-initiated nocturnal recesses, 56% were accompanied by evidence of depredation at the nest during the subsequent nest monitoring visit. Hen-initiated nocturnal recesses began later in the night (closer to morning) and were shorter than predator-initiated nocturnal recesses. Our results indicate that nocturnal incubation recesses occur regularly (14% of all recesses) and, similar to diurnal recesses, most nocturnal recesses (75%) are initiated by the hen rather than an approaching predator.

Interrupted incubation: How dabbling ducks respond when flushed from the nest.

Nesting birds must provide a thermal environment sufficient for egg development while also meeting self-maintenance needs. Many birds, particularly those with uniparental incubation, achieve this balance through periodic incubation recesses, during which foraging and other self-maintenance activities can occur. However, incubating birds may experience disturbances such as predator or human activity which interrupt natural incubation patterns by compelling them to leave the nest. We characterized incubating mallard Anas platyrhynchos and gadwall Mareca strepera hens' responses when flushed by predators and investigators in Suisun Marsh, California, USA. Diurnal incubation recesses initiated by investigators approaching nests were 63% longer than natural diurnal incubation recesses initiated by the hen (geometric mean: 226.77 min versus 142.04 min). Nocturnal incubation recesses, many of which were likely the result of predators flushing hens, were of similar duration regardless of whether the nest was partially depredated during the event (115.33 [101.01;131.68] minutes) or not (119.62 [111.96;127.82] minutes), yet were 16% shorter than natural diurnal incubation recesses. Hens moved further from the nest during natural diurnal recesses or investigator-initiated recesses than during nocturnal recesses, and the proportion of hen locations recorded in wetland versus upland habitat during recesses varied with recess type (model-predicted means: natural diurnal recess 0.77; investigator-initiated recess 0.82; nocturnal recess 0.31). Hens were more likely to take a natural recess following an investigator-initiated recess earlier that same day than following a natural recess earlier that same day, and natural recesses that followed an investigator-initiated recess were longer than natural recesses that followed an earlier natural recess, suggesting that hens may not fulfill all of their physiological needs during investigator-initiated recesses. We found no evidence that the duration of investigator-initiated recesses was influenced by repeated visits to the nest, whether by predators or by investigators, and trapping and handling the hen did not affect investigator-initiated recess duration unless the hen was also fitted with a backpack-harness style GPS-GSM transmitter at the time of capture. Hens that were captured and fitted with GPS-GSM transmitters took recesses that were 26% longer than recesses during which a hen was captured but a GPS-GSM transmitter was not attached. Incubation interruptions had measurable but limited and specific effects on hen behavior.

Timing, frequency, and duration of incubation recesses in dabbling ducks.

Nest attendance is an important determinant of avian reproductive success, and identifying factors that influence the frequency and duration of incubation recesses furthers our understanding of how incubating birds balance their needs with those of their offspring. We characterized the frequency and timing (start time, end time, and duration) of incubation recesses for mallard (Anas platyrhynchos) and gadwall (Mareca strepera) hens breeding in Suisun Marsh, California, USA, and examined the influences of day of year, ambient temperature at the nest, incubation day, and clutch size on recess frequency and timing using linear mixed models. Mallard, on average, took more recesses per day (1.69 ± 0.80, mean ± standard deviation) than did gadwall (1.39 ± 0.69), and 45% of mallard nest-days were characterized by two recesses, while only 27% of gadwall nest-days were characterized by two recesses. Mallard morning recesses started at 06:14 ± 02:46 and lasted 106.11 ± 2.01 min, whereas mallard afternoon recesses started at 16:39 ± 02:11 and lasted 155.39 ± 1.99 min. Gadwall morning recesses started at 06:30 ± 02:46 and lasted 91.28 ± 2.32 min, and gadwall afternoon recesses started at 16:31 ± 01:57 and lasted 192.69 ± 1.89 min. Mallard and gadwall started recesses earlier in the day with increasing ambient temperature, but later in the day as the season progressed. Recess duration decreased as the season progressed and as clutch size increased, and increased with ambient temperature at the nest. The impending darkness of sunset appeared to be a strong cue for ending a recess and returning to the nest, because hens returned to their nests earlier than expected when recesses were expected to end after sunset. Within hens, the timing of incubation recesses was repeatable across incubation days and was most repeatable for mallard afternoon recesses and on days in which hens took only one recess. Hens were most likely to be away from nests between 04:00 and 07:00 and between 16:00 and 19:00; therefore, investigators should search for nests between 07:00 and 16:00. Our analyses identified important factors influencing incubation recess timing in dabbling ducks and have important implications for nest monitoring programs.

Mercury exposure in mammalian mesopredators inhabiting a brackish marsh.

Bioaccumulation of environmental contaminants in mammalian predators can serve as an indicator of ecosystem health. We examined mercury concentrations of raccoons (Procyon lotor; n = 37 individuals) and striped skunks (Mephitis mephitis; n = 87 individuals) in Suisun Marsh, California, a large brackish marsh that is characterized by contiguous tracts of tidal marsh and seasonally impounded wetlands. Mean (standard error; range) total mercury concentrations in adult hair grown from 2015 to 2018 were 28.50 µg/g dw (3.05 µg/g dw; range: 4.46-81.01 µg/g dw) in raccoons and 4.85 µg/g dw (0.54 µg/g dw; range: 1.53-27.02 µg/g dw) in striped skunks. We reviewed mammalian hair mercury concentrations in the literature and raccoon mercury concentrations in Suisun Marsh were among the highest observed for wild mammals. Although striped skunk hair mercury concentrations were 83% lower than raccoons, they were higher than proposed background levels for mercury in mesopredator hair (1-5 µg/g). Hair mercury concentrations in skunks and raccoons were not related to animal size, but mercury concentrations were higher in skunks in poorer body condition. Large inter-annual differences in hair mercury concentrations suggest that methylmercury exposure to mammalian predators varied among years. Mercury concentrations of raccoon hair grown in 2017 were 2.7 times greater than hair grown in 2015, 1.7 times greater than hair grown in 2016, and 1.6 times greater than hair grown in 2018. Annual mean raccoon and skunk hair mercury concentrations increased with wetland habitat area. Furthermore, during 2017, raccoon hair mercury concentrations increased with the proportion of raccoon home ranges that was wetted habitat, as quantified using global positioning system (GPS) collars. The elevated mercury concentrations we observed in raccoons and skunks suggest that other wildlife at similar or higher trophic positions may also be exposed to elevated methylmercury bioaccumulation in brackish marshes.

Sitting ducklings: Timing of hatch, nest departure, and predation risk for dabbling duck broods.

For ground-nesting waterfowl, the timing of egg hatch and duckling departure from the nest may be influenced by the risk of predation at the nest and en route to wetlands and constrained by the time required for ducklings to imprint on the hen and be physically able to leave the nest. We determined the timing of hatch, nest departure, and predation on dabbling duck broods using small video cameras placed at the nests of mallard (Anas platyrhynchos; n = 26), gadwall (Mareca strepera; n = 24), and cinnamon teal (Anas cyanoptera; n = 5). Mallard eggs began to hatch throughout the day and night, whereas gadwall eggs generally started to hatch during daylight hours (mean 7.5 hr after dawn). Among all species, duckling departure from the nest occurred during daylight (98%), and 53% of hens typically left the nest with their broods 1-4 hr after dawn. For mallard and gadwall, we identified three strategies for the timing of nest departure: (a) 9% of broods left the nest the same day that eggs began to hatch (6-12 hr later), (b) 81% of broods left the nest the day after eggs began to hatch, and (c) 10% of broods waited 2 days to depart the nest after eggs began to hatch, leaving the nest just after the second dawn (27-42 hr later). Overall, eggs were depredated at 10% of nests with cameras in the 2 days prior to hatch and ducklings were depredated at 15% of nests with cameras before leaving the nest. Our results suggest that broods prefer to depart the nest early in the morning, which may best balance developmental constraints with predation risk both at the nest and en route to wetlands.

GPS tracking data reveals daily spatio-temporal movement patterns of waterfowl.

BACKGROUND: Spatio-temporal patterns of movement can characterize relationships between organisms and their surroundings, and address gaps in our understanding of species ecology, activity budgets, bioenergetics, and habitat resource management. Highly mobile waterfowl, which can exploit resources over large spatial extents, are excellent models to understand relationships between movements and resource usage, landscape interactions and specific habitat needs. METHODS: We tracked 3 species of dabbling ducks with GPS-GSM transmitters in 2015-17 to examine fine-scale movement patterns over 24 h periods (30 min interval), dividing movement pathways into temporally continuous segments and spatially contiguous patches. We quantified distances moved, area used and time allocated across the day, using linear and generalized linear mixed models. We investigated behavior through relationships between these variables. RESULTS: Movements and space-use were small, and varied by species, sex and season. Gadwall (Mareca strepera) generally moved least (FFDs: 0.5-0.7 km), but their larger foraging patches resulted from longer within-area movements. Pintails (Anas acuta) moved most, were more likely to conduct flights > 300 m, had FFDs of 0.8-1.1 km, used more segments and patches per day that they revisited more frequently, resulting in the longest daily total movements. Females and males differed only during the post-hunt season when females moved more. 23.6% of track segments were short duration (1-2 locations), approximately 1/3 more than would be expected if they occurred randomly, and were more dispersed in the landscape than longer segments. Distance moved in 30 min shortened as segment duration increased, likely reflecting phases of non-movement captured within segments. CONCLUSIONS: Pacific Flyway ducks spend the majority of time using smaller foraging and resting areas than expected or previously reported, implying that foraging areas may be highly localized, and nutrients obtainable from smaller areas. Additionally, movement reductions over time demonstrates behavioral adjustments that represent divergent energetic demands, the detection of which is a key advantage of higher frequency data. Ducks likely use less energy for movement than currently predicted and management, including distribution and configuration of essential habitat, may require reconsideration. Our study illustrates how fine-scale movement data from tracking help understand and inform various other fields of research.