Body mass regulation during incubation in female common eiders Somateria mollissima

We investigated changes in incubation behaviour induced by body fuel depletion in incubating female common eiders, which, in contrast to pelagic seabirds, fast despite being close to marine food sources. In the Svalbard Archipelago, electronic scales were placed under eider nests and the incubation of six birds was prolonged by using wax-filled eggs. Based on changes in the rate of body mass loss in normally incubating females and in ten captive birds that did not incubate, body reserves neared depletion on average four days after hatching. During prolonged incubation, females took more frequent and longer recesses. Nest attentiveness consequently decreased, but was still high. In contrast to recesses during normal incubation, during which body mass of the birds decreased, mass remained constant during the recesses of prolonged incubation. The body stores of female eiders seemed to enable them to complete incubation with a limited safety margin. A further drop in body mass is avoided when a critical body mass is reached, because birds then start feeding enough to maintain mass while continuing incubation. Presumably, a similar mechanism will enable eiders to continue incubation when body reserves are prematurely depleted before hatching.     

Do extended incubation recesses carry fitness costs in two cavity‐nesting birds?

Because extended incubation recesses, where incubating songbirds are away from nests for periods much longer than usual, occur infrequently, they have been treated as outliers in most previous studies and thus overlooked. However, egg temperatures can potentially fall below the physiological zero temperature during extended recesses, potentially affecting developing embryos. As such, evaluating extended recesses in an ecological context and identifying their possible fitness effects are important. With this aim, we used iButton data loggers to monitor the incubation behavior of female Blue Tits (Cyanistes caeruleus) and Great Tits (Parus major) during two breeding seasons in central Spain. We classified incubation recesses as extended if they were more than four times the mean recess duration for each species. Extended incubation recesses occurred more frequently in 2012 when females exhibited poorer body condition. Female Blue Tits had more extended incubation recesses than female Great Tits and, for both species, more extended recesses occurred at the beginning of the breeding season. Both nest attentiveness and average minimum nest temperature decreased when at least one extended recess occurred. Incubation periods averaged 4 d longer for nests where females had at least one extended recess, potentially increasing predation risk and resulting in lower‐quality nestlings. Overall, our results suggest that extended recesses may be more common among songbirds than previously thought and that, due to their effects on egg temperatures and attentiveness, they could impose fitness costs.     

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.     

Incubation recess behaviors influence nest survival of Wild Turkeys

In ground nesting upland birds, reproductive activities contribute to elevated predation risk, so females presumably use multiple strategies to ensure nest success. Identification of drivers reducing predation risk has primarily focused on evaluating vegetative conditions at nest sites, but behavioral decisions manifested through movements during incubation may be additional drivers of nest survival. However, our understanding of how movements during incubation impact nest survival is limited for most ground nesting birds. Using GPS data collected from female Eastern Wild Turkeys (n = 206), we evaluated nest survival as it relates to movement behaviors during incubation, including recess frequency, distance traveled during recesses, and habitat selection during recess movements. We identified 9,361 movements off nests and 6,529 recess events based on approximately 62,065 hr of incubation data, and estimated mean nest attentiveness of 84.0%. The numbers of recesses taken daily were variable across females (range: 1?7). Nest survival modeling indicated that increased cumulative distance moved during recesses each day was the primary driver of positive daily nest survival. Our results suggest behavioral decisions are influencing trade‐offs between nest survival and adult female survival during incubation to reduce predation risk, specifically through adjustments to distances traveled during recesses.     

Time allocation between feeding and incubation in uniparental arctic-breeding shorebirds: energy reserves provide leeway in a tight schedule

Birds with uniparental incubation may face a time allocation problem between incubation and feeding. Eggs need regular warming to hatch successfully, but the parent must leave the nest to feed and safeguard its own survival. Time allocation during incubation is likely to depend on factors influencing egg cooling rates, parental energy requirements and feeding intake rate. How this allocation problem is resolved was subject of this study on arctic‐breeding shorebirds. We compared incubation rhythms between four uniparental shorebird species differing in size and expected to find both species differences and weather effects on the organisation of incubation. Attentive behaviour and responses to variation in weather showed a remarkable consistency across species. All species alternated feeding bouts (recesses) with brooding bouts throughout the day. Recesses were concentrated in the warmer parts of the day, while recess duration showed little diurnal variation. Despite continuous daylight, a pronounced day‐night rhythmicity was apparent. The four species in this study spent a similar proportion (13–19%) of the time off their nest. After correction for weather effects, the number of recesses was largest in the smallest species, while recess duration was longest in the largest species. Total recess time per day increased on cold days through an increase of mean recess length, while the number of recesses decreased. Comparing our observations to predictions derived from criteria that birds might use to organise their attentive behaviour, showed that the limits are set by parental requirements, while the energy stores of adults provide some leeway for short‐term adjustments to environmental variability. If breeding birds trade off feeding time against incubation time, energy stores are expected to be influenced by weather. We expected uniparental species to be more likely to show weather effects on condition than biparentals, as in the latter ‘off duty’ time is much larger and independent of weather. This prediction was tested by comparing energy stores in two uniparental species and a biparental congener. While body mass of uniparental incubators decreased after a period with low temperatures, body mass of the biparental species did not.     

State-dependent incubation behaviour in the high arctic barnacle geese

Energetic trade-offs between time spent on incubation and times spent on foraging for nesting birds give individuals in good body condition the possibility to incubate more continuously. In the present paper, the incubation behaviour of female barnacle geese Branta leucopsis was quantified in a colony in Svalbard. Females were weighted in early incubation and feeding recess lengths and frequencies were recorded. The feeding behaviour during the course of incubation was significantly correlated to by body mass, and heavy females had both fewer and shorter feeding recesses than lighter females. Moreover, there was an increase in the number of feeding recesses and the total feeding time as the incubation period progressed. Neither clutch size nor egg laying date had an effect on incubation behaviour. However, clutch size was positively related to female body mass suggesting that high-quality females produce large clutches but also allocate more body reserves to incubation. Females left the nest to feed at all times of the day, but more frequently during day time. This was not related to their body mass. Females presumably leave their nest at the time of day when the costs to reheat eggs are at a minimum. The diurnal rhythm may also be adjusted to the activity by egg predators. Overall the results support the state-dependent hypothesis for incubation behaviour, suggesting that body condition at the start of incubation is an important factor for incubation behaviour in barnacle geese.     

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.     

Incubation behavior of king eiders on the coastal plain of northern Alaska

Incubating birds balance their energetic demands during incubation with the needs of the developing embryos. Incubation behavior is correlated with body size; larger birds can accumulate more endogenous reserves and maintain higher incubation constancy. King eiders (Somateria spectabilis) contend with variable and cold spring weather, little nesting cover, and low food availability, and thus are likely to rely heavily on endogenous reserves to maintain high incubation constancy. We examined the patterns of nest attendance of king eiders at Teshekpuk and Kuparuk, Alaska (2002–2005) in relation to clutch size, daily temperature, and endogenous reserves to explore factors controlling incubation behavior. Females at Kuparuk had higher constancy (98.5 ± 0.2%, n = 30) than at Teshekpuk (96.9 ± 0.8%, n = 26), largely due to length of recesses. Mean recess length ranged from 21.5 to 23.7 min at Kuparuk, and from 28.5 to 51.2 min at Teshekpuk. Mean body mass on arrival at breeding grounds (range; Teshekpuk 1,541–1,805, Kuparuk 1,616–1,760), and at the end of incubation (Teshekpuk 1,113–1,174, Kuparuk 1,173–1,183), did not vary between sites or among years (F < 1.1, P > 0.3). Daily constancy increased 1% with every 5°C increase in minimum daily temperature (β _min = 0.005, 95% CI 0.002, 0.009). Higher constancy combined with similar mass loss at Kuparuk implies that females there met foraging requirements with shorter recesses. Additionally, females took more recesses at low temperatures, suggesting increased maintenance needs which were potentially ameliorated by feeding during these recesses, indicating that metabolic costs and local foraging conditions drove incubation behavior.     

Extended incubation recesses by alpine‐breeding Horned Larks: a strategy for dealing with inclement weather?

ABSTRACT Incubating birds can incur high energetic costs and, when faced with a trade‐off between incubation and foraging, parents may neglect their eggs in favor of their own somatic needs. Extended incubation recesses are an example of neglect, but they are often treated as outliers and largely overlooked in studies of incubation behavior. We studied incubation rhythms of Horned Larks (Eremophila alpestris) on Hudson Bay Mountain, British Columbia, Canada, during four breeding seasons. Incubation recesses averaged 10.92 ± 0.38 min (N= 4076 2‐h periods), but we observed 70 extended recesses, ranging from 59 to 387 min in duration, at 35 nests. Although rare (<1% of all daytime recesses), extended recesses occurred in all 4 yr, were longer and more frequent in colder years (60% occurred in the two coldest years), and often occurred during inclement weather (39% occurred during three storm events). Extended recesses did not appear to compensate for long attendance periods because extended recess duration was not correlated with the duration of previous on‐bouts (P= 0.10, N= 70) or the mean on‐bout duration of the previous 2‐h period (P= 0.36, N= 70). Rather, extended recesses seemed to reflect a shift in parental investment away from their eggs and toward self‐maintenance when faced with energetically stressful conditions. Extended recesses may have reduced embryo viability; egg‐hatching rates were 91 ± 2.4% for nests where females did not take extended recesses and 81 ± 4.2% for nests where females did take extended recesses (P= 0.02, N= 56 nests). Extended recesses during incubation are rare events, but they may represent an important mechanism that allows birds to breed successfully in energetically challenging conditions.     

Movements of female Sage Grouse Centrocercus urophasianus during incubation recess

We combined GPS data‐loggers, VHF transmitters and DVR video‐monitoring to measure fine‐scale movement patterns during daily incubation recesses by female Sage Grouse Centrocercus urophasianus, a species with uniparental incubation that has experienced widespread population decline and distributional contraction. Most (69.6%) Sage Grouse recess activity was highly localized within a core recess area averaging 2.58 ± 0.64 ha, and females remained within 242.3 ± 30.0 m from the nest during recesses (total recess areas were 11.06 ± 2.27 ha). Visually conspicuous Sage Grouse movements near nests at the start and end of recesses and consistent occupation of core recess areas point to a mechanism for newly abundant predators such as the Northern Raven Corvus corax to detect and depredate Sage Grouse nests. Our methods apply to other avian species of scientific interest and conservation concern.     

Effects of implanted satellite transmitters on behavior and survival of female common eiders

Surgically implanted satellite transmitters have been widely used in studies of avian ecology, yet little is known about their potential impacts on birds. We implanted satellite transmitters with percutaneous antennae (approx. 50 g) in 17 female common eiders (Somateria mollissima) at a breeding colony in Arctic Canada. Among females implanted during incubation, 11 of 12 nests were abandoned within 1 week of being radioed. We observed no differences in the proportion of time that implanted female eiders allocated to basic behaviors. Radioed birds were more likely to pick or preen their abdominal (site of surgical incision) and posterior–dorsal (site of antenna exit) regions than unmarked females, although these behaviors were rare (approx. 0.3% of total time budget). Three of 10 females re-observed had a pronounced limp following surgery, but we observed no walking difficulties among these females in subsequent seasons, and we observed some implanted eiders nesting in subsequent years. Mark–resighting models suggest eiders with transmitters had lower apparent survival the year after implantation (67.0%; 85% CI: 47.8–81.9%) than did color-banded eiders (87.5%, 85% CI: 82.5–91.2%), but there was no model support for a survival difference in subsequent years. We conclude that transmitter implantation in common eiders leads to short-term changes in behavior and a decline in first year survival. We encourage researchers to collect similar data on their study subjects where possible and use it to determine the degree to which data are representative of the greater population. © 2011 The Wildlife Society.     

Use of the Beaufort Sea by King Eiders Breeding on the North Slope of Alaska

Abstract: We estimated areas used by king eiders (Somateria spectabilis) in the Alaskan Beaufort Sea, how distributions of used areas varied, and characteristics that explained variation in the number of days spent at sea, to provide regulatory agencies with baseline data needed to minimize impacts of potential offshore oil development. We implanted sixty king eiders with satellite transmitters at nesting areas on the North Slope of Alaska, USA, in 2002-2004. More than 80% of marked eiders spent >2 weeks staging offshore prior to beginning a postbreeding molt migration. During postbreeding staging and migration, male king eiders had much broader distributions in the Alaskan Beaufort Sea than female eiders, which were concentrated in Harrison and Smith Bays. Distribution did not vary by sex during spring migration in the year after marking. Shorter residence times of eiders and deeper water at locations used during spring migration suggest the Alaskan Beaufort Sea might not be as critical a staging area for king eiders during prebreeding as it is postbreeding. Residence time in the Beaufort Sea varied by sex, with female king eiders spending more days at sea than males in spring and during postbreeding. We conclude the Alaskan Beaufort Sea is an important staging area for king eiders during postbreeding, and eider distribution should be considered by managers when mitigating for future offshore development. We recommend future studies examine the importance of spring staging areas outside the Alaskan Beaufort Sea.     

Foraging time and dietary intake by breeding Ross's and Lesser Snow Geese

We compared foraging times of female Ross's (Chen rossii) and Lesser Snow Geese (Chen caerulescens caerulescens) breeding at Karrak Lake, NT, Canada and examined variation due to time of day and reproductive stage. We subsequently collected female geese that had foraged for known duration and we estimated mass of foods consumed during foraging bouts. Female Ross's Geese spent more time foraging (mean % - SE =28.4ǃ.3%; P=0.0002), on average, than did female Lesser Snow Geese (21.5 - 1.4%). Foraging time by female geese differed among reproductive stages, but differences were not consistent among time periods (stage-by-time block interaction, P=0.0003). Females spent considerably more time foraging during prelaying and laying than during incubation. Ross's Geese also spent a greater percent of time feeding (83.0DŽ.8%) during incubation recesses than did Lesser Snow Geese (60.9Dž.6%). Consumption of organic matter during foraging bouts was minimal; estimated consumption averaged 9.6dž.0 and 12.4dž.6 g (mean - SE) dry mass/day before incubation and 5.9DŽ.0 and 5.7DŽ.1 g dry mass/day during incubation for Lesser Snow and Ross's Geese, respectively. Diets consisted primarily of mosses (bryophytes), Chickweed (Stellaria spp.) and Sedges (Carex spp.). Before incubation, eggshell consumption was estimated as 4.3Dž.2 and 0.4ǂ.3 g dry mass/day for Lesser Snow and Ross's Geese, respectively; neither species consumed eggshell during incubation. We conclude that eggshell from nests of previous years is likely an important source of dietary calcium used to meet mineral demands of eggshell formation at Karrak Lake. Our findings of wide disparities between foraging time and food intake indicate that results from studies that do not directly measure intake rates remain equivocal. Finally, we propose four hypotheses accounting for foraging effort that evidently yields little nutritional or energetic benefit to geese nesting at Karrak Lake.     

Night conditions affect morning incubation behaviour differently across a latitudinal gradient

Intermittently incubating birds alternate between sessions of egg warming and recesses for foraging during the day, but stay on the nest continuously at night. Hence, energy costs of nocturnal incubation (which increase during longer and colder nights) cannot be replenished until the next day. Night conditions might therefore be expected to affect morning incubation behaviour the day after. We tested this prediction by exploring latitudinal and seasonal trends in characteristics of the first recess in Eastern Bluebirds Sialia sialis over a 1400-km latitudinal gradient in the continental USA. The time from civil dawn to leaving the nest (latency) increased with latitude early in the breeding season but decreased with latitude late in the season. Birds breeding at higher latitudes also took longer first recesses throughout the season, which led to a larger drop in nest temperature. At the local scale, birds rose earlier after longer nights if the night was also cold, but night length did not predict latency following warm nights. The first recess was longer if the night was warmer, probably because birds could replenish reserves at lower risk of low egg temperature. Our study shows that characteristics of the night led to behavioural changes in features of early morning incubation that were evident at both continental and local scales. These responses also affected nest temperature. Hence, night conditions carry over to incubation behaviour the following morning, which in turn may impose thermal constraints on embryonic development.     

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.     

Effects of down collection on incubation temperature, nesting behaviour and hatching success of common eiders (Somateria mollissima) in west Iceland

Incubating common eiders (Somateria mollissima) insulate their nests with down to maintain desirable heat and humidity for their eggs. Eiderdown has been collected by Icelandic farmers for centuries, and down is replaced by hay during collection. This study determined whether down collecting affected the female eiders or their hatching success. We compared the following variables between down and hay nests: incubation temperature in the nest, incubation constancy, recess frequency, recess duration, egg rotation and hatching success of the clutch. Temperature data loggers recorded nest temperatures from 3 June to 9 July 2006 in nests insulated with down (n = 12) and hay (n = 12). The mean incubation temperatures, 31.5 and 30.7°C, in down and hay nests, or the maximum and minimum temperatures, did not differ between nest types where hatching succeeded. Cooling rates in down, on average 0.34°C/min and hay nests 0.44°C/min, were similar during incubation recesses. Females left their nests 0–4 times every 24 h regardless of nest type, for a mean duration of 45 and 47.5 min in down and hay nests, respectively. The mean frequency of egg rotation, 13.9 and 15.3 times every 24 h, was similar between down and hay nests, respectively. Hatching success adjusted for clutch size was similar, 0.60 and 0.67 in down and hay nests. These findings indicate that nest down is not a critical factor for the incubating eider. Because of high effect sizes for cooling rate and hatching success, we hesitate to conclude that absolutely no effects exist. However, we conclude that delaying down collection until just before eggs hatch will minimize any possible effect of down collection on females.     

Factors influencing Cinnamon Teal nest attendance patterns

Patterns of nest attendance in birds result from complex behaviours and influence the success of reproductive events. Incubation behaviours vary based on individual body condition, energy requirements and environmental factors. We assessed nest attendance patterns in Cinnamon Teal Spatula cyanoptera breeding in the San Luis Valley of Colorado in 2016–2017 using trail and video cameras to observe behaviours throughout incubation. We evaluated the effect of temporal, life‐history and environmental covariates on the frequency and duration of incubation recesses as well as the incubation constancy. There was considerable model uncertainty among the models used to evaluate recess frequency. Recess duration varied according to the interaction between nest age and a quadratic effect of time of day, with hens on older nests taking longer recesses in the afternoon and hens on nests earlier in incubation taking longer recesses in the morning and evening. Incubation constancy decreased with higher ambient temperatures in the study area. This study provides evidence that Cinnamon Teal modify their behaviour during incubation according to the age of the nest and the time of day. These results improve our knowledge of Cinnamon Teal breeding ecology and shed light on the behaviours that fast‐lived species may use to cope with environmental factors during nesting.     

Activity budgets of a marked common loon (Gavia immer) nesting population

A newly devised nightlighting technique was used to capture breeding adult common loons (Gavia immer) at the Seney National Wildlife Refuge in northern Michigan in 1989. The behaviors of 6 pairs of known-sex, color-marked common loons were subsequently quantified during the breeding cycle in 1990. Collected observational data indicate that foraging, resting, locomotion, and preening were frequent throughout the breeding cycle. Time spent foraging was greatest during the pre-nesting period (53 to 57%), but declined significantly during the nesting and post-nesting periods (p<0.05). Time spent foraging during the pre-nesting period was similar to that of fall and winter studies. During the pre-nesting period adult loons spent about 15% of the time in locomotion; this was significantly greater than the other time periods (p<0.05) and is attributed to selecting a nest site. During the nesting cycle, almost half of each bird's time spent was nest-sitting. Sexual differences were negligible during nest-sitting. Resting and chick-rearing were the predominant behaviors during the post-nesting period and were responsible for the biggest difference in parental duties. Time spent preening declined from 8% during the pre-nesting period to 4 to 5% during the post-nesting period. Time spent by nesting pairs to produce chicks is approximately 10% during pre-nesting, 48 to 49% during nesting, and between 38 to 44% during post-nesting. By quantifying and establishing behavioral standards, subtle abnormalities or changes can be detected to better manage for viable common loon populations.     

Low incubation investment in the burrow‐nesting Crab Plover Dromas ardeola permits extended foraging on a tidal food resource

We used GPS data‐loggers, video‐recordings and dummy eggs to assess whether foraging needs may force the low incubation attentiveness (< 55%) of the Crab Plover Dromas ardeola, a crab‐eating wader of the Indian Ocean that nests colonially in burrows. The tidal cycle was the major determinant of the time budget and some foraging trips were more distant from the colony than previously known (up to 26 km away and lasting up to 45 h). The longest trips were mostly made by off‐duty parents, but on‐duty parents also frequently left the nest unattended while foraging for 1–7 h. However, the time spent at the colony area (47%) and the time spent roosting on the foraging grounds (16%) would have allowed almost continuous incubation, as in other species with shared incubation. Therefore, the low incubation attentiveness is not explained by the need for long foraging trips but is largely dependent on a high intermittent rhythm of incubation with many short recesses (5.8 ± 2.6 recesses/h) that were not spent foraging but just outside the burrow or thermoregulating at the seashore. As a result, the eggs were warmed on average only 1.7 °C above burrow temperature, slightly more during high tide periods and when burrow temperature was lower between 20:00 and 10:00 h, only partly counteracting the temperature fluctuations of the incubation chamber. These results suggest that low incubation attentiveness is due to the favourable thermal conditions provided by safe nesting burrows and by the hot tropical breeding season, a combination that allows simultaneous foraging by parents and the exploitation of distant foraging grounds. Why Crab Plovers engage in many short recesses from incubation still remains to be clarified but the need to thermoregulate at the seashore and to watch for predators may play a role.     

Do T3 levels in incubating eiders reflect the cost of incubation among clutch sizes?

Complete development of avian eggs requires external heat, inducing in most species an energetic cost of incubation for the parents. Triiodothyronine (T(3)) has been implicated in the control of the metabolic rate and is decreased during fasting in most bird species. This raises the question of the regulation of T(3) during reproduction when incubation (thus heat production) is associated with fasting (and energy sparing). In this study, plasma concentrations of T(3) were studied for different clutch sizes in incubating, as well as in nonincubating, fasting female eiders. Our results show that the T(3) levels decrease during fasting in nonincubating birds, whereas they were maintained during the incubation fast. T(3) levels increased in female eiders at hatching. The plasma T(3) level did not vary among natural clutch sizes in eiders but did so when manipulated. T(3) levels increased when eggs were added (to a maximum of six eggs, i.e., the biggest natural clutch size) or removed (to two eggs, i.e., the smallest natural clutch size). Our results suggest that (1) high T(3) levels during incubation may participate to a threshold of heat production and incubation metabolic rate in eiders despite the fact that they are fasting; (2) since T(3) is associated with the energy expenditure in birds, incubating an enlarged or reduced clutch size may lead to a higher energetic cost of incubation in eiders; and (3) the energy demand of the ducklings at hatching is probably important, as the female T(3) concentrations are then at their highest levels. Thus, any modification of the natural clutch size leads to a rise in the T(3) level of the incubating female, suggesting an additional cost of incubation. Knowing that there is no variation of T(3) levels among natural clutch sizes, this study suggests that a female eider produces a number of eggs corresponding to the energy she can invest in incubation.