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.

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.

Which egg features predict egg rejection responses in American robins? Replicating Rothstein's (1982) study.

Rothstein (Behavioral Ecology and Sociobiology, 11, 1982, 229) was one of the first comprehensive studies to examine how different egg features influence egg rejection behaviors of avian brood parasite-hosts. The methods and conclusions of Rothstein (1982) laid the foundation for subsequent experimental brood parasitism studies over the past thirty years, but its results have never been evaluated with replication. Here, we partially replicated Rothstein's (1982) experiments using parallel artificial model egg treatments to simulate cowbird (Molothrus ater) parasitism in American robin (Turdus migratorius) nests. We compared our data with those of Rothstein (1982) and confirmed most of its original findings: (1) robins reject model eggs that differ from the appearance of a natural robin egg toward that of a natural cowbird egg in background color, size, and maculation; (2) rejection responses were best predicted by model egg background color; and (3) model eggs differing by two or more features from natural robin eggs were more likely to be rejected than model eggs differing by one feature alone. In contrast with Rothstein's (1982) conclusion that American robin egg recognition is not specifically tuned toward rejection of brown-headed cowbird eggs, we argue that our results and those of other recent studies of robin egg rejection suggest a discrimination bias toward rejection of cowbird eggs. Future work on egg recognition will benefit from utilizing a range of model eggs varying continuously in background color, maculation patterning, and size in combination with avian visual modeling, rather than using model eggs which vary only discretely.

Using 3D printed eggs to examine the egg-rejection behaviour of wild birds.

The coevolutionary relationships between brood parasites and their hosts are often studied by examining the egg rejection behaviour of host species using artificial eggs. However, the traditional methods for producing artificial eggs out of plasticine, plastic, wood, or plaster-of-Paris are laborious, imprecise, and prone to human error. As an alternative, 3D printing may reduce human error, enable more precise manipulation of egg size and shape, and provide a more accurate and replicable protocol for generating artificial stimuli than traditional methods. However, the usefulness of 3D printing technology for egg rejection research remains to be tested. Here, we applied 3D printing technology to the extensively studied egg rejection behaviour of American robins, Turdus migratorius. Eggs of the robin's brood parasites, brown-headed cowbirds, Molothrus ater, vary greatly in size and shape, but it is unknown whether host egg rejection decisions differ across this gradient of natural variation. We printed artificial eggs that encompass the natural range of shapes and sizes of cowbird eggs, painted them to resemble either robin or cowbird egg colour, and used them to artificially parasitize nests of breeding wild robins. In line with previous studies, we show that robins accept mimetically coloured and reject non-mimetically coloured artificial eggs. Although we found no evidence that subtle differences in parasitic egg size or shape affect robins' rejection decisions, 3D printing will provide an opportunity for more extensive experimentation on the potential biological or evolutionary significance of size and shape variation of foreign eggs in rejection decisions. We provide a detailed protocol for generating 3D printed eggs using either personal 3D printers or commercial printing services, and highlight additional potential future applications for this technology in the study of egg rejection.

Potential Mechanisms Driving Population Variation in Spatial Memory and the Hippocampus in Food-caching Chickadees.

Harsh environments and severe winters have been hypothesized to favor improvement of the cognitive abilities necessary for successful foraging. Geographic variation in winter climate, then, is likely associated with differences in selection pressures on cognitive ability, which could lead to evolutionary changes in cognition and its neural mechanisms, assuming that variation in these traits is heritable. Here, we focus on two species of food-caching chickadees (genus Poecile), which rely on stored food for survival over winter and require the use of spatial memory to recover their stores. These species also exhibit extensive climate-related population level variation in spatial memory and the hippocampus, including volume, the total number and size of neurons, and adults' rates of neurogenesis. Such variation could be driven by several mechanisms within the context of natural selection, including independent, population-specific selection (local adaptation), environment experience-based plasticity, developmental differences, and/or epigenetic differences. Extensive data on cognition, brain morphology, and behavior in multiple populations of these two species of chickadees along longitudinal, latitudinal, and elevational gradients in winter climate are most consistent with the hypothesis that natural selection drives the evolution of local adaptations associated with spatial memory differences among populations. Conversely, there is little support for the hypotheses that environment-induced plasticity or developmental differences are the main causes of population differences across climatic gradients. Available data on epigenetic modifications of memory ability are also inconsistent with the observed patterns of population variation, with birds living in more stressful and harsher environments having better spatial memory associated with a larger hippocampus and a larger number of hippocampal neurons. Overall, the existing data are most consistent with the hypothesis that highly predictable differences in winter climate drive the evolution and maintenance of differences among populations both in cognition and in the brain via local adaptations, at least in food-caching parids.

Experimental shifts in intraclutch egg color variation do not affect egg rejection in a host of a non-egg-mimetic avian brood parasite.

Avian brood parasites lay their eggs in the nests of other birds, and impose the costs associated with rearing parasitic young onto these hosts. Many hosts of brood parasites defend against parasitism by removing foreign eggs from the nest. In systems where parasitic eggs mimic host eggs in coloration and patterning, extensive intraclutch variation in egg appearances may impair the host's ability to recognize and reject parasitic eggs, but experimental investigation of this effect has produced conflicting results. The cognitive mechanism by which hosts recognize parasitic eggs may vary across brood parasite hosts, and this may explain variation in experimental outcome across studies investigating egg rejection in hosts of egg-mimicking brood parasites. In contrast, for hosts of non-egg-mimetic parasites, intraclutch egg color variation is not predicted to co-vary with foreign egg rejection, irrespective of cognitive mechanism. Here we tested for effects of intraclutch egg color variation in a host of nonmimetic brood parasite by manipulating egg color in American robins (Turdus migratorius), hosts of brown-headed cowbirds (Molothrus ater). We recorded robins' behavioral responses to simulated cowbird parasitism in nests where color variation was artificially enhanced or reduced. We also quantified egg color variation within and between unmanipulated robin clutches as perceived by robins themselves using spectrophotometric measures and avian visual modeling. In unmanipulated nests, egg color varied more between than within robin clutches. As predicted, however, manipulation of color variation did not affect rejection rates. Overall, our results best support the scenario wherein egg rejection is the outcome of selective pressure by a nonmimetic brood parasite, because robins are efficient rejecters of foreign eggs, irrespective of the color variation within their own clutch.

The role of egg-nest contrast in the rejection of brood parasitic eggs.

Hosts of avian brood parasites can avoid the reproductive costs of raising genetically unrelated offspring by rejecting parasitic eggs. The perceptual cues and controls mediating parasitic egg discrimination and ejection are well studied: hosts are thought to use differences in egg color, brightness, maculation, size and shape to discriminate between their own and foreign eggs. Most theories of brood parasitism implicitly assume that the primary criteria to which hosts attend when discriminating eggs are differences between the eggs themselves. However, this assumption is confounded by the degree to which chromatic and achromatic characteristics of the nest lining co-vary with egg coloration, so that egg-nest contrast per se might be the recognition cue driving parasitic egg detection. Here, we systematically tested whether and how egg-nest contrast itself contributes to foreign egg discrimination. In an artificial parasitism experiment, we independently manipulated egg color and nest lining color of the egg-ejector American robin (Turdus migratorius), a host of the obligate brood parasitic brown-headed cowbird (Molothrus ater). We hypothesized that the degree of contrast between foreign eggs and the nest background would affect host egg rejection behavior. We predicted that experimentally decreasing egg-nest chromatic and achromatic contrast (i.e. rendering parasitic eggs more cryptic against the nest lining) would decrease rejection rates, while increasing egg-nest contrast would increase rejection rates. In contrast to our predictions, egg-nest contrast was not a significant predictor of egg ejection patterns. Instead, egg color significantly predicted responses to parasitism. We conclude that egg-egg differences are the primary drivers of egg rejection in this system. Future studies should test for the effects of egg-nest contrast per se in predicting parasitic egg recognition in other host-parasite systems, including those hosts building enclosed nests and those parasites laying cryptic eggs, as an alternative to hypothesized effects of egg-egg contrast.