Incubation and brooding rhythm of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Incubation and brooding performance of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, in the 1991 -1992 austral summer is reported in detail and compared with data from two other areas. There was an inverse relationship between the mean foraging trip length and mean peak weight of chicks. Variation in shift length throughout incubation resulted from a combination of seasonal effects (food supply) and behavioural adjustment to hatching. The length of the last incubation shift was independent of its number but decreased with date of initiation. The average weight of nestattending birds increased steadily during the incubation period and at the same rate in both sexes. Females attained seasonal peak body-weight before the egg hatched, whereas males just recovered their pre-breeding weight. Males took on a slightly larger share of incubation (52%, range 40–63%) and invested more time in mutual nest attendance. Length of foraging trips varied consistently in pairs and individuals, its repeatability being twice as high in females as in males. The incubation performance of individual females (length of the first and an average foraging trip) was correlated with the size of the egg they had laid and also to subsequent chick growth and fledging success. There was a tendency in successful pairs to spend more time together at the nest and to have shorter foraging trips. I suggest that variation in breeding performance among pairs was mainly a result of individual variation in female quality, with pairs where the female contributed more to incubation being more successful. There is little evidence that egg losses were caused by parental errors.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

Incubation Feeding and Nest Attentiveness in a Socially Monogamous Songbird: Role of Feather Colouration,Territory Quality and Ambient Environment

Parental investment and environmental conditions determine reproductive success in wild‐ranging animals. Parental effort during incubation, and consequently factors driving it, has profound consequences for reproductive success in birds. The female nutrition hypothesis states that high male feeding enables the incubating female to spend more time on eggs, which can lead to higher hatching success. Moreover, both male and female parental investment during incubation might be signalled by plumage colouration. To test these hypotheses, we investigated relationships between male and female incubation behaviour and carotenoid and melanin‐based plumage colouration, territory quality and ambient temperature in the Great Tit Parus major. We also studied the effect of female incubation behaviour on hatching success. Intensity of male incubation feeding increased with lower temperatures and was higher in territories with more food supply, but only in poor years with low overall food supply. Female nest attentiveness increased with lower temperatures. Plumage colouration did not predict incubation behaviour of either parent. Thus, incubation behaviour of both parents was related mainly to environmental conditions. Moreover, there was no relationship between male incubation feeding, female nest attentiveness and hatching success. Consequently, our data were not consistent with the female nutrition hypothesis.     

Breeding strategies of Antarctic Petrels Thalassoica antarctica and Southern Fulmars Fulmarus glacialoides in the high Antarctic and implications for reproductive success

Breeding strategies of two closely related fulmarine petrels were studied on Ardery Island, on the continental coast of East Antarctica, where short summers are expected to narrow the time-window for reproduction. Both species had a similar breeding period (97 days from laying to fledging) but Antarctic Petrels Thalassoica antarctica bred up to 16 days earlier than Southern Fulmars. During the pre-laying exodus, all Antarctic Petrels deserted the colony, whereas some Southern Fulmars Fulmarus glacialoides remained. Antarctic Petrels exhibited stronger synchronization in breeding, made longer foraging trips and spent less time guarding their chicks than Southern Fulmars. Overall breeding success of both species was similar but failures of Antarctic Petrels were concentrated in the early egg-phase and after hatching, when parents ceased guarding. Southern Fulmars lost eggs and chicks later in the breeding cycle and so wasted more parental investment in failed breeding attempts. Different breeding strategies may be imposed by flight characteristics; Southern Fulmars are less capable of crossing large expanses of pack ice and need to delay breeding until the sea ice retreats and breaks up. However, due to the short summer they risk chick failure when weather conditions deteriorate late in the season.     

Weight loss of Antarctic Fulmars Fulmarus glacialoides during incubation and chick brooding

Weight losses incurred by incubating birds of both sexes, and weight changes at the beginning and end of shifts between successive incubation and brooding shifts were studied in the Antarctic Fulmar Fulmarus glacialoides in Adélie Land, Antarctica. The overall rate of weight loss during incubation fasts was 4.46% per day in both sexes but 3.91% per day after the first 2 days of fast once the stomach contents had been digested. Antarctic Fulmars appear to have rates of body-weight loss significantly greater than the rates of weight loss observed in any other species of albatross or petrel. The validity of methods using the rate of weight loss to estimate incubation metabolic rates is discussed, and it appears that for birds incubating for periods of less than 2–3 days the digestion of the stomach contents can bias the estimation. The absence of change in weight at the beginning of shifts during incubation and brooding, together with the short time spent on foraging trips, suggests that food availability is high and/or food is readily accessible to Antarctic Fulmars. The availability of food allows the Antarctic Fulmar to compensate for the high energy cost of breeding that could be due to the cold Antarctic environment and perhaps also to a higher intrinsic metabolism.     

Sensitivity of breeding parameters to food supply in Black-legged Kittiwakes Rissa tridactyla

We fed Herring Clupea pallasi to pairs of Black-legged Kittiwakes Rissa tridactyla throughout the breeding season in two years at a colony in the northern Gulf of Alaska. We measured responses to supplemental feeding in a wide array of breeding parameters to gauge their relative sensitivity to food supply, and thus their potential as indicators of natural foraging conditions. Conventional measures of success (hatching, fledging and overall productivity) were more effective as indicators of food supply than behavioural attributes such as courtship feeding, chick provisioning rates and sibling aggression. However, behaviour such as nest relief during incubation and adult attendance with older chicks were also highly responsive to supplemental food and may be useful for monitoring environmental conditions in studies of shorter duration. On average, the chick-rearing stage contained more sensitive indicators of food availability than prelaying or incubation stages. Overall, rates of hatching and fledging success, and the mean duration of incubation shifts were the most food-sensitive parameters studied.     

Incubation scheduling by Northern Fulmars ( Fulmarus glacialis ) in the Canadian High Arctic

Northern Fulmars (Fulmarus glacialis) are a ubiquitous seabird of the North Atlantic and North Pacific oceans that breed in colonies surrounded by markedly different marine environmental conditions. I have studied the incubation behavior of fulmars at a remote colony in the Canadian High Arctic, where the birds had to cross 200 km of sea-ice during the early incubation period to reach feeding areas. These fulmars completed incubation in fewer shifts and had a longer mean incubation shift duration (5.3 days) than their counterparts breeding in the Boreal oceanographic zone. In particular, the mean duration of the first incubation shift by males in successful pairs (10.3 days) was 2 days longer than that for males in unsuccessful pairs and was longer than that reported at any other colony. This exceptionally long shift by the male may be required at this site to give females enough time to recoup energetic reserves after egg-laying, at a time when marine productivity in the Arctic is still seasonably low.     

Incubation patterns in a central-place forager affect lifetime reproductive success: scaling of patterns from a foraging bout to a lifetime

Background

Long-lived seabirds face a conflict between current and lifelong reproductive success. During incubation shifts, egg neglect is sometimes necessary to avoid starvation, but may compromise the current reproductive attempt. However, factors underlying this decision process are poorly understood. We focus on the ancient murrelet, Synthliboramphus antiquus, an alcid with exceptionally long incubation shift lengths, and test the impact of environmental factors on incubation shift length in relation to reproductive success.

Methodology/Principal Findings

Using an information theoretic approach, we show that incubation shift length was a strong predictor of reproductive success for ancient murrelets at Reef Island, Haida Gwaii, British Columbia, Canada during the 2007 and 2008 breeding seasons. The most important factors explaining an individual''s shift length were egg size, wind speed and the length of the mate''s previous shift. Wind speed and tide height were the two most important factors for determining foraging behavior, as measured by dive frequency and depth.

Conclusions/Significance

Our study demonstrates that (i) species-specific reproductive strategies interact with environmental conditions such as wind speed to form multiple incubation patterns and (ii) maintaining regular incubation shifts is an essential component of reproductive success.     

The influence of food supply on the breeding ecology of Kittiwakes Rissa tridactyla in Shetland

We measured the breeding performance, body condition, time budgets and foraging ranges of Kittiwakes Rissa tridactyla at Sumburgh Head, Shetland, in two years of contrasting food availability. Kittiwakes in Shetland generally feed their young almost entirely on sandeels, and fisheries data indicated that stocks of sandeels in Shetland waters were at least ten times higher in 1991 than in 1990. Fledging success of Kittiwakes was nil in 1990 and 68% of eggs laid in 1991, although clutch-size and hatching success were no different between years. Post-hatching foraging trips in 1991 were of comparable duration to those recorded at other colonies in conditions of good food supply (2–3 h), while trips recorded during incubation or post-hatching in 1990 were approximately three times longer on average than at corresponding stages of the breeding season in 1991. Radio-tracking data indicated that adults generally stayed within 5 km of the colony in 1991 but flew more than 40 km from the colony on each trip in 1990. Eggs were apparently not left unattended in either year, despite the fact that this required adults to incubate for periods in excess of 44 h in 1990. The extent to which adults were able to increase trip durations, foraging ranges and incubation shift lengths between years, while maintaining hatching success, indicates the degree to which Kittiwakes are normally buffered against adverse feeding conditions during incubation. Reduced nest attendance and lower body-condition of adults post-hatching in 1990, in conjunction with complete post-hatching breeding failure, indicate that adults were beyond the limits of their buffering capacity during chick-rearing in 1990.     

Incubation routines and foraging-trip regulation in the Grey-faced Petrel Pterodroma macroptera gouldi

In Grey-faced Petrels Pterodroma macroptera gouldi parental care during the 55-day incubation period is mainly achieved by three long shifts (two by the male) when one partner is ashore while the other forages at sea. This situation would seem to carry high risks of parental desertion and failure of the hatching chick to receive a meal before its yolk sac is exhausted. We examined incubation routines of 51 pairs of Grey-faced Petrels to investigate factors implicated in parental desertion and in the duration of the final foraging trip before hatching. The duration of the final long foraging trip (by the female) during incubation is not predicted by the time spent fasting on the nest but by the time elapsed since laying. Desertions accounted for 6.5% of egg losses. Eggs were deserted not after the delay of the foraging bird to return to the nest, but when the incubating bird departed early in the incubation spell. Desertions were not a response to critically depleted energy reserves.     

Incubation feeding as a male tactic for early hatching

Male marsh tits, Parus palustris, regularly feed their mates from the beginning of nest building until hatching. Over three periods (the 15 days preceding egg formation, egg formation/laying and incubation) the number of food passes by the male to the female increased significantly. There was a significant negative relationship between the frequency with which the male fed the female in the nest during incubation and the length of the incubation period. Female blue tits, Parus caeruleus, experimentally supplied with food in the nestbox during incubation had a significantly shorter incubation period than control females. Clutches of experimentally fed females also tended to hatch more successfully. It is concluded that feeding of the female by the male is a nutritional contribution and that the shorter incubation period and increased hatching success enhance the fitness of both parents. However, the male should balance the benefits against the costs in time and energy and therefore not necessarily work at a maximal level. In accordance with this is the finding that the male's provisioning rate increased when ambient temperatures decreased. Adverse weather may jeopardize the whole or large proportions of the clutch, thereby significantly reducing the benefit from the current breeding attempt.     

Influence of incubation temperature on hatching success,energy expenditure for embryonic development,and size and morphology of hatchlings in the oriental garden lizard,Calotes versicolor (Agamidae)

We incubated eggs of Calotes versicolor at four constant temperatures ranging from 24 degrees C to 33 degrees C to assess the effects of incubation temperature on hatching success, embryonic use of energy, and hatchling phenotypes that are likely to affect fitness. All viable eggs increased in mass throughout incubation due to absorption of water, and mass gain during incubation was dependent on initial egg mass and incubation temperature. The average duration of incubation at 24 degrees C, 27 degrees C, 30 degrees C, and 33 degrees C was 82.1 days, 60.5 days, 51.4 days, and 50.3 days, respectively. Incubation temperature affected hatching success, energy expenditure for embryonic development, and several hatchling traits examined, but it did not affect the sex ratio of hatchlings. Hatching success was lowest (3.4%) at 33 degrees C, but a higher incidence of deformed embryos was recorded from eggs incubated at this temperature compared to eggs incubated at lower temperatures. Most of the deformed embryos died at the last stage of incubation. Energy expenditure for embryonic development was, however, higher in eggs incubated at 33 degrees C than those similarly incubated at lower temperatures. A prolonged exposure of eggs of C. versicolor at 33 degrees C appears to have an adverse and presumably lethal effect on embryonic development. Hatching success at 24 degrees C was also low (43.3%), but hatchlings incubated at 24 degrees C did not differ in any of the examined traits from those incubated at two intermediate temperatures (27 degrees C and 30 degrees C). Hatchlings incubated at 33 degrees C were smaller (snout-vent length, SVL) than those incubated at lower incubation temperatures and had larger mass residuals (from the regression on SVL) as well as shorter head length, hindlimb length, tympanum diameter, and eye diameter relative to SVL. Hatchlings from 33 degrees C had significantly lower scores on the first axis of a principal component analysis representing mainly SVL-free head size (length and width) and fore- and hindlimb lengths, but they had significantly higher scores on the second axis mainly representing SVL-free wet body mass. Variation in the level of fluctuating asymmetry in eye diameter associated with incubation temperatures was quite high, and it was clearly consistent with the prediction that environmental stress associated with the highest incubation temperatures might produce the highest level of asymmetry. Newly emerged hatchlings exhibited sexual dimorphism in head width, with male hatchlings having larger head width than females.     

Incubation feeding by male Scarlet Tanagers: a mate removal experiment

ABSTRACT Incubation feeding, where males feed their mates, is a common behavior in birds and may improve female condition, nest attentiveness, and nesting success. We used behavioral observations and a temporary mate removal experiment to test the female nutrition hypothesis for incubation feeding by male Scarlet Tanagers (Piranga olivacea). All males (N= 20) were observed incubation feeding and fed females both at the nest (x? 1.36 trips/h) and away from the nest (x? 20.1 trips/h). Male feeding rate off‐nest was negatively correlated with the duration of female foraging bouts and positively correlated with the total time females spent incubating per hour. Eggs were predated at seven of 19 (37%) nests, but nest survival during incubation was not related to either female incubation behavior or male feeding rate. During temporary removal experiments (N= 12), female Scarlet Tanagers remained on the nest significantly longer and did not have longer foraging bouts. An unexpected outcome of the removal experiments was a dramatic change in female vocal behavior. All 12 experimental females gave chik‐burr calls during the male‐removal experiments (x? bout length = 11.7 min), but during normal observation periods only six of 20 females at the incubation stage gave chik‐burr calls (x? bout length = 0.7 min, N= 20). Our results suggest that female tanagers likely gain nutritional benefits from incubation feeding, but male feeding may not improve immediate reproductive success. Nine of 54 (17%) nestlings in five of 17 broods (29%) were extra‐pair young (EPY), indicating that males could potentially benefit from incubation feeding via mate retention and fidelity as well as, or instead of, through immediate gains in reproductive success. Our study indicates that females benefit from incubation feeding and do not simply passively accept food from their mates, but instead may influence male feeding rates through direct (e.g., mate following and vocalizing) and indirect (the threat of mate abandonment or cuckoldry) means.     

Mate-sharing costs in polygynous Northern Lapwings Vanellus vanellus

In this study bigamous female Northern Lapwings Vanellus vanellus received significantly less incubation relief from their males than monogamous females. On average, monogamous males spent 34.3% of their time incubating and bigamous males 29.9%. Bigamous males divided their effort between their nests, incubating on average 9.4% on primary nests and 20.5% on secondary nests. Bigamous females compensated for the lack of male relief. Primary females incubated for 71.8% of their time, secondary females for 64.2%, while monogamous females spent 52.7% of their time incubating. As a result, there was no significant difference in total nest attentiveness among nests of different status. Primary and secondary females received equivalent incubation relief from the male. Bigamous males increased their contribution to incubation significantly as the season progressed. A bigamous male's distribution of incubation relief between his females was unrelated to female body mass, or to the degree of asynchrony between primary and secondary females in arrival and laying. Incubation time was significantly, negatively, correlated with total nest attentiveness. Monogamous females spent most time, secondary females spent an intermediate time, and primary females spent the least time on maintenance behaviour (foraging, comfort behaviour, inactivity). No significant differences were found in hatching success among females of different mating status. However, the ratio of unhatched to hatched eggs (i.e. the eggs that remained in the nest at the time of hatching) differed significantly: secondary females hatched a smaller proportion of their eggs than monogamous and primary females.     

Time and energy budgets of breeding males and females in sandgrouse Pterocles species

The investment of time and energy in reproduction by male and female sandgrouse (genus Pterocles ) was studied. Energy expenditure was estimated using time budgets and laboratory measurements of metabolism. Female parental expenditure was greatest during laying and incubation. Females took the larger share of daytime incubation, which reduced their time available for foraging (3.6 h compared with 4.8 h for males). To remain in energy balance, they required a higher food intake rate than males (97 kJ/h compared with 79 kJ/h, respectively) and consequently were likely to lose condition during incubation. Male parental expenditure was greatest during chick-rearing, especially when the young were well grown but still dependent on the male for water. The role of males as water carriers increased their energy expenditure and decreased their time available for foraging. Males also took the greater share of time spent in vigilance on behalf of the young, which further reduced their foraging time (3.2 h compared with 4.9 h for females). To remain in energy balance, males with young required a higher intake rate than females (62 kJ/h compared with 44 kJ/h, respectively).     

Size-assortative Monogamy and Paternal Egg Care in a Coral Goby Paragobiodon echinocephalus

Paragobiodon echinocephalus (Pisces; Gobiidae) inhabits the branching coral Stylophora pistillata. Only the largest two fish bred in a coral irrespective of group size, which was larger in larger corals. Breeding pairs consisted of a male and female of similar body size. Eggs were spawned on the coral branch and guarded by the male parent for about 4 days until hatching. Successive spawning occurred usually one day after hatching. The reproductive success of a pair, measured as the number of newly hatched larvae, was positively correlated with the body size of the male as well as the female. That is, not only female fecundity but also male ability of egg guarding were limited by body size almost equally. This should promote size-assortative monogamy in the goby.     

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.     

Regulation of foraging trips and incubation routine in male and female wandering albatrosses

The resolution of the conflict between eggcare and foraging was studied in male and female wandering albatrosses. The foraging zone and range, duration of incubation shifts and foraging trips, and associated changes in body mass were studied. Costs during incubation, expressed as the time spent incubating and the proportional loss of body mass, were similar for both sexes. The mass gained at sea was related to the duration of foraging trips, but the relationship was much less significant in males, where foraging ranges, though similar on average to those of females, were very variable. Males foraged in more southerly waters than females, and gained mass more rapidly. Only females appeared to regulate the duration of foraging trips, and this compensated for the mass lost during the incubation fast. Previous breeding experience had no influence on foraging efficiency. Egg desertion because of depletion of body reserves was very rare because birds have a wide safety margin, i.e. the difference between the average body mass when relieved and that at nest desertion. This safety margin enables the birds to compensate for the high variability in the duration of foraging trips, and is probably a reason for the high breeding success of wandering albatrosses. Decisions to return from the sea to the nest or to desert the nest are probably related to the status of body reserves, and have been selected in the large wandering albatross so that both present and future reproductive success are maximised.     

Parental effects on fertilization and hatching success and development of Atlantic halibut (Hippoglossus hippoglossus L.) embryos and larvae

The parental effects on fertilization and early life history traits were studied in Atlantic halibut, Hippoglossus hippoglossus L. Sperm from 12 different males were used to fertilize eggs of two females in separate crosses. The fertilization success were generally high, above 80% of developing embryos at 16-cell stage in 20 of 24 crosses with an average of 85.9+/-17.6% and 87.2+/-16.5% for female A and female B, respectively. Corresponding hatching success was 74.8+/-17.7% and 41.6+/-20.1%, respectively. The relationship between fertilization success and hatching success was positive. The parental influence on hatching was dominated by a strong and significant (p<0.001) maternal effect; however, the paternal effect was also significant (p<0.001). Furthermore, there was no relationship between fertilization success, hatching success and larvae viability as a high number of larvae developed locked jaws (i.e., were not functional). There was a significant (p<0.01) difference in yield of functional larvae of female A (43%) and female B (56%), and also between crosses sired by different males. The standard length of offspring of female A (12.4+/-0.5 mm) and B (12.6+/-0.6 mm) were significantly (p<0.001) different, and also significantly influenced by both the female (p<0.001) and the male (p<0.001). The present paper provides strong indications that not only the female, but also the male parent influences quantitative features of early development of their offspring.     

Egg variability and hatching success in the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Eggs of the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, exhibited high variability in volume between females (up to 48%), while for each individual female, both volume and shape of eggs were highly correlated over two seasons. Both female size and body-condition at laying were unrelated to egg-volume in the 'good'season 1991, but larger females produced larger eggs in the 'bad'season 1990, when eggs were smaller on average. In 1991, females in better body-condition, but not larger, started laying earlier. In spite of synchronized laying (75% of eggs laid within five days), egg-volume decreased with laying date. Egg-volume differed significantly between the two years, which could not be explained by changes in the breeding population or timing of breeding. The laying of relatively small eggs and low overall breeding success in 1990 most probably reflected changes in food availability. Overall hatchability was 88% and did not differ significantly between disturbed and control colonies in 1991. Larger and more rounded eggs showed better hatching success, but hatchability was related more to egg-volume whereas hatching rate was related more to egg-shape. As the majority of egg losses were attributable to predation, the hatching rate was influenced by parental performance relatively more than by hatchability. Hence the relationship between hatching rate and egg-shape probably reflects improvement in hatching success with female age/experience, whereas the relationship between egg-size and hatchability suggests effect of egg-size independently of parental traits.