Nest attendance by tropical and temperate passerine birds: Same constancy,different strategy

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Energetics of small body size and high latitude the Rufous Hummingbird in coastal Alaska

The Rufous Hummingbird is the smallest bird breeding at latitudes of 58° to 60°. Nesting chronology and food sequence was observed. The microclimate of the nest-site and its potential for heat loss during incubation were similar to those of hummingbirds in the Rocky Mountains at 39°N. Hummingbirds of lower latitudes increase their active-daylength in direct proportion to solar daylength (active day = 1.03 × solar day), whereas the Rufous Hummingbird tended to utilize less of the long daylength of coastal Alaska (active day = 197 min + 0.83 × solar day).     

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.     

BREEDING BIOLOGY OF THE BEARDED VULTURE IN SOUTHERN AFRICA,PART I: THE PRELAYING AND INCUBATION PERIODS

Brown, C. J. 1990. Breeding biolo of the Bearded Vulture in southern Africa, Part I: The pre-laying and incubation periods. Ostrich 61: 24–32.

In southern Africa the Bearded Vulture Gpaetus barbatus lays its eggs in mid-winter. between the second half of May and the first week of July. Pairs became more active in their nesting areas about six weeks before laying and usually roosted there at night. Courtship flights were less frequent and demonstrative than in Eurasian birds and took place mainly in the late afternoons. During the pre-laying period most nest visits (77%) were to bring nesting material, 92% by the male. All nesting material was arranged by the female. Copulation was always preceded by allopreening, and occurred most frequently in the mornings. No copulation or courtship display took place after the first egg had been laid. Of 18 clutches, 16 (89%) contained two eggs and the remainder one egg. The laying interval was usually 3–5 days (range 2–9 days). Incubation started with the first egg and was evenly shared by both parents during the day, but only the female incubated at night, individual pairs maintained distinctive nest attendance and foraging period timetables, which allowed sufficient time for self-foraging by both parentes. No food was brought into the nest during the pre-laying and incubation periods, but in some pairs food was cached in nearby potholes in cliffs. The incubation period was 56–57 days.     

Partial incubation during egg laying reduces eggshell microbial loads in a temperate‐breeding passerine

Incubation prior to clutch completion may be adaptive if it maintains egg viability by inhibiting eggshell microbial growth, thus reducing the likelihood that the embryo becomes infected. To test this hypothesis, we examined the effect of partial incubation during egg laying on eggshell microbial loads in eastern bluebirds Sialia sialis breeding at a temperate‐zone site. We sampled eggshell microbes prior to and following four days of exposure to either partial incubation during the laying period or ambient environmental conditions without incubation (experimental eggs). Microbial colony counts declined significantly for eggs left in the nest during the laying period but did not vary significantly for eggs exposed to ambient conditions. Initial microbial loads were more similar to those previously reported from tropical than temperate environments, and microbes from potentially pathogenic groups were detected on 88% of first‐laid eggs on the day of laying. Egg viability was maintained when eggs were held indoors for four days without incubation but declined sharply thereafter. Our results suggest that partial incubation during egg laying may enhance egg viability in eastern bluebirds by reducing eggshell microbial loads; these effects appear stronger than those usually reported from the temperate zone.     

Trade-offs in nest site selection in coastal populations of Lapwings Vanellus vanellus

In coastal populations of Lapwings Vanellus vanellus in southwestern Sweden, arable fields predominated as foraging habitat before laying. Females caught more large prey items on arable fields and shores than on pastures. Close to egg laying, females foraged mainly near their future nest sites. Arable land and pastures were used to a similar extent for nesting. We found no difference in nest predation between habitats. Egg volume varied among females and was correlated with wing-length, body mass and condition. Mean egg volume also was positively correlated with feeding time on arable land before laying. Pairs nesting on arable fields therefore generally produced larger eggs than those on pastures. The distances between nests and chick foraging areas, however, were significantly longer for birds nesting on arable land than for those on pastures. Moreover, in 2 of 3 years, the proportion of hatched chicks that survived until fledging was negatively correlated with this distance. There was no difference in chick survival between broods hatched on arable fields and pastures. We suggest that nest site selection and offspring production involve a trade-off between the benefits of nesting close to rich feeding grounds for adults and the costs of moving long distances between nest sites and chick-rearing areas.     

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

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

Relationship of weight loss to ambient humidity of birds eggs during incubation

Summary Eggs of birds nesting in wet and dry habitats, have been artificially incubated at controlled humidity white weight loss of the eggs and shell water vapour conductance have been determined. Eggs of species from wet habitats loose weight at a higher rate than those from drier habitats at a given relative humidity.It is suggested that the conductance of the egg shell to water vapour is adapted to the conditions of humidity in the environment such that weight loss varies little (and less than predictable) in relation to the relative humidity at the nesting sites.The relative humidity surrounding eggs during natural incubation was found to be in the range of 30–50% in 4 different species. Humidity in the nest during natural incubation was found to be higher than what would result if ambient air was heated to incubation temperature indicating that the sitting bird conserves humidity around the eggs.     

How avian incubation behaviour influences egg surface temperatures: relationships with egg position,development and clutch size

While understanding heat exchange between incubating adults and their eggs is central to the study of avian incubation energetics, current theory based on thermal measurements from dummy eggs reveals little about the mechanisms of this heat exchange or behavioural implications for the incubating bird. For example, we know little about how birds distribute their eggs based on temperature differences among egg positions within the nest cup. We studied the great tit Parus major, a species with a large clutch size, to investigate surface cooling rates of individual eggs within the nest cup across a range of ambient temperatures in a field situation. Using state‐of‐the‐art portable infrared imaging and digital photography we tested for associations between egg surface temperature (and rate of cooling) and a combination of egg specific (mass, shape, laying order, position within clutch) and incubation specific (clutch size, ambient temperature, day of incubation) variables. Egg surface temperature and cooling rates were related to the position of the eggs within the nest cup, with outer eggs being initially colder and cooling quicker than central eggs. Between foraging bouts, females moved outer eggs significantly more than centrally positioned eggs. Our results demonstrate that females are capable of responding to individual egg temperature by moving eggs around the nest cup, and that the energy cost to the female may increase as incubation proceeds. In addition, our results showing that smaller clutches experience lower initial incubation temperatures and cool quicker than larger clutches warrant further attention for optimal clutch size theory and studies of energetic constraints during incubation. Finally, researchers using dummy eggs to record egg temperature have ignored important elements of contact‐incubation, namely the complexity of how eggs cool and how females respond to these changes.     

Mating success in brood-guarding male rock bass,Ambloplites rupestris: the effect of body size

Synopsis The hypothesis that variation in a paternal trait associated with offspring survival will result in female mate choice based on that trait was tested in rock bass,Ambloplites rupestris, a temperate fresh water fish with uniparental male care. The number of eggs acquired by 108 nesting male rock bass, in Cranberry Lake (New York State, U.S.A.), was estimated in two different spawning episodes that differed in the size structure of the nesting male stock. Early breeders (between the 4–7 June 1989) were mainly medium and large males (101–300 g), as compared to late breeders (between the 19–21 June 1989) which were mostly small (50–100 g). Because survival of larvae in rock bass is significantly lower in nests guarded by small males as compared to nests guarded by medium and large males, it was hypothesized that female mate choice should have been (1) size-based, and (2) more intense late in the season when the nesting stock was comprised mostly of small males. The results were consistent with these hypotheses. Variation in egg numbers per nest was significantly higher during the late spawning episode. In addition, egg number per nest were normally distributed during the early spawning episode, but were positively skewed and leptokurtic during the later episode. The correlation between male size and egg number was significant for both spawning episodes. However, during the late episode male size explained 23% of the variation in egg numbers versus only 8% of the variation during the earlier episode.     

Interspecific variation in desiccation survival time of Aedes (Stegomyia) mosquito eggs is correlated with habitat and egg size

Summary Survival times of eggs under three humidity conditions (42%, 68%, 88% RH) were investigated among Aedes (Stegomyia) mosquitoes from temperate and tropical zones (5 species and 20 geographical strains). This subgenus tends to occupy small aquatic sites as larvae, where desiccation resistance of eggs is necessary during habitat drought. Interspecific comparison showed that the egg survival time was correlated with egg volume and dryness of source locality, and probably with habitat. Aedes aegypti is associated most with arid climate and human-disturbed habitats — its large eggs survived the longest periods at all humidities. Aedes albopictus ranges from tropics to temperate zones and inhabits both disturbed and forest habitats — its eggs were less desiccation-resistant than A. aegypti eggs. The survival times for forest species eggs (A. riversi, A. galloisi, A. flavopictus) were variable at high humidities but at the lowest humidity were consistently shorter than for eggs of A. aegypti and A. albopictus.     

Western Snowy Plovers Charadrius alexandrinus nivosus select nesting substrates that enhance egg crypsis and improve nest survival

Predation is an important cause of nest failure for many birds and has shaped the life‐history characteristics of many species, especially ground‐nesting shorebirds. We examined nesting success, causes of clutch failure and nest survival in relation to variation in substrate characteristics in a colour‐marked population of Western Snowy Plovers Charadrius alexandrinus nivosus breeding on riverine gravel bars in coastal northern California. Plovers experienced higher nesting success on gravel bars than on nearby beaches, which were characterized by more homogeneous, sandy substrates. On gravel bars, Plovers nested in habitats characterized by large, heterogeneous substrates, with more egg‐sized stones, compared with random sites. Egg crypsis, as indexed by time required of a naïve observer to detect a nest, increased with number of egg‐sized substrates. Nest survival correlated negatively with heterogeneity of substrates and positively with the number of egg‐sized stones. Consistently high nesting success of Plovers on gravel bars indicates that this high‐quality habitat deserves special management considerations given the species’ threatened status.     

Male boobies expel eggs when paternity is in doubt

We analyzed the effect of increased risk of cuckoldry on maleparental investment in eggs in the colonial blue-footed booby(Sula nebouxii). Seventeen experimental males were removedfrom the nesting territory for 10-12 h on a single day, 1-5days before laying (females' supposed fertile period = SFP),and 17 control males were removed for the same amount of timeon a single day, 7-29 days before laying (before the SFP).These removals were intended to simulate extended absence fromthe nest on a foraging excursion. Female extrapair courtshipand copulation rates did not increase during the removal ofthe social mate, and there was no evidence that experimentaland control males differed quantitatively in incubation ordefense of the clutch. However, 43% of experimental males expelledthe first-laid egg from the nest, whereas no control male didso. Apparently, male boobies drastically reduce parental investmentin eggs with a presumed elevated probability of extrapair fertilizationby destroying them.     

Intraguild predation between spawning smallmouth bass (Micropterus dolomieu) and nest‐raiding crayfish (Orconectes rusticus): implications for bass nesting success

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Do alien predators pose a particular risk to duck nests in Northern Europe? Results from an artificial nest experiment

Several alien predator species have spread widely in Europe during the last five decades and pose a potential enhanced risk to native nesting ducks and their eggs. Because predation is an important factor limiting Northern Hemisphere duck nest survival, we ask the question, do alien species increase the nest loss risk to ground nesting ducks? We created 418 artificial duck nests in low densities around inland waters in Finland and Denmark during 2017–2019 and monitored them for seven days after construction using wildlife cameras to record whether alien species visit and prey on the nests more often than native species. We sampled various duck breeding habitats from eutrophic agricultural lakes and wetlands to oligotrophic lakes and urban environments. The results differed between habitats and the two countries, which likely reflect the local population densities of the predator species. The raccoon dog (Nyctereutes procyonoides), an alien species, was the most common mammalian nest visitor in all habitats and its occurrence reduced nest survival. Only in wetland habitats was the native red fox (Vulpes vulpes) an equally common nest visitor, where another alien species, the American mink (Neovison vison), also occurred among nest visitors. Although cautious about concluding too much from visitations to artificial nests, these results imply that duck breeding habitats in Northern Europe already support abundant and effective alien nest predators, whose relative frequency of visitation to artificial nests suggest that they potentially add to the nest predation risk to ducks over native predators.

     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

Multi-scale habitat selection and foraging ecology of the eurasian hoopoe (<Emphasis Type="Italic">Upupa epops</Emphasis>) in pine plantations

Bird conservation can be challenging in landscapes with high habitat turnover such as planted forests, especially for species that require large home ranges and juxtaposition of different habitats to complete their life cycle. The eurasian hoopoe (Upupa epops) has declined severely in western Europe but is still abundant in south-western France. We studied habitat selection of hoopoes in pine plantation forests using a multi-scale survey, including point-counts at the landscape level and radio-tracking at the home-range scale. We quantified habitat use by systematically observing bird behaviour and characterized foraging sites according to micro-habitat variables and abundance of the main prey in the study area, the pine processionary moth (Thaumetopoea pityocampa). At the landscape scale, hoopoes selected habitat mosaics of high diversity, including deciduous woods and hedgerows as main nesting sites. At the home-range scale, hoopoes showed strong selection for short grassland vegetation along sand tracks as main foraging habitats. Vegetation was significantly shorter and sparser at foraging sites than random, and foraging intensity appeared to be significantly correlated with moth winter nest abundance. Hoopoe nesting success decreased during the three study years in line with processionary moth abundance. Thus, we suggest that hoopoes need complementation between foraging and breeding habitats to establish successfully in pine plantations. Hoopoe conservation requires the maintenance of adjacent breeding (deciduous woods) and foraging habitats (short swards adjacent to plantation edges), and consequently depends on the maintenance of habitat diversity at the landscape scale.     

Revisiting the influence of aggressive interactions on the survival of the first‐laid egg in crested penguins (genus Eudyptes)

Crested penguins Eudyptes spp. have evolved a unique form of breeding in which the first of two eggs laid is much smaller than the second and has a higher likelihood of being lost during egg laying and incubation. In this study, we quantified aggressive behaviour in nesting Snares penguins and undertook an egg survival analysis to examine which factors influence egg loss. During 120 h of observation of 50 nests, we recorded a total of 300 aggressive events in which females were repeatedly pecked, bitten and beaten. Aggressive events lasted from less than a minute to up to 55 min (mean 4.6 ± 7.4 min). Single males were the aggressor in 75% of aggressive events and in 50.7% of aggressive events the aggressor was identified as a neighbouring, breeding male. A greater percentage of the small first eggs (34%) were lost than the large second eggs (4%). We found that egg mortality was influenced by 1) whether the other egg within a nest had hatched, 2) who was present at the nest (father, mother or both) and 3) the average duration of aggressive events on the nest. When one egg within a nest had hatched, the other egg had a vastly increased mortality risk irrespective of aggression. However, long, aggressive events directed towards females after their partners had gone foraging, also increased the probability of egg loss. We suggest that the prolonged nest attendance by breeding males well beyond egg laying is in response to the high frequency of aggressive behaviour during this time. Our data show that A‐egg losses occur due to intraspecific aggression in this species. Further research is needed to clarify whether aggressive behaviour in breeding crested penguins is modulated by elevated testosterone levels in the males and whether any reproductive benefits accrue to the aggressors.