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.     

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.     

Parental care behavior in the monogamous,sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

I studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti-predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.     

DIET,HOME RANGE,HUNTING AND REPRODUCTIVE BEHAVIOUR OF A PAIR OF DICKINSON'S KESTREL FALCO DZCKZNSONZ IN THE KRUGER NATIONAL PARK,SOUTH AFRICA

Summary

BENN, G.A. &; KEMP, A.C. 1995. Diet, home range, hunting and reproductive behaviour of a pair of Dickinson's Kestrel Falco dickinsoni in the Kruger National Park, South Africa. Ostrich 66: 81–91.

During July-December 1992, the diet, home range, hunting and reproductive behaviour of a pair of Dickinson's Kestrel Falco dickinsoni was recorded in the Kruger National Park, South Africa. Numerically, for both sexes combined, invertebrates formed the majority (56%) of the diet, while separately the female caught 75% and the male 49% invertebrate prey. During courtship and incubation the male supplied the female with primarily vertebrate prey and both provisioned mainly vertebrates to the nestlings (male = 80%; female = 57%). The non-breeding home range of the female was 27.8 km², and the breeding home range of the male was 26.3 km2. Both utilised their home ranges differentially, the area within a 2 km radius of the nest (12.6 km²) being used proportionally more than the remaining area. The home range of the female was compared to that of other Falco spp. and was larger than would be expected based on body weight. Perch-hunting was the only technique utilised by both sexes, with 79–80% of observed strike attempts from dead trees. During the day, the 9.emale spent 87% and the male 77% of the time perch-hunting, with respective hunting success rates of 69% and 58%. During courtship, the female spent much of her time (94%) close to the nest, where the male supplied her with prey. During incubation, the male spent 95% of his time within 2 km of the nest tree, where he hunted to supply the female with prey at a rate of 0.3 items.hr^?1 and assisted in nest defence. On occasion the male entered the nest to relieve the female, and remained in the cavity on average for 134 min (n = 5). As the young got older, the female spent less time at the nest and provisioned more items to the nestlings. Overall, there was an increase in the rate of prey provisioning to the nestlings from 0.45 items.hr^?1 (10 days old) to 0.85 items.hr^?1 (21 days old). The male initially passed prey to the female but provisioned directly to older nestlings.     

Temporal changes in prey composition and biomass delivery to African Crowned Eagle nestlings in urban areas of KwaZulu-Natal,South Africa

Globally urban areas are expanding rapidly and this usually has negative effects on biodiversity. Despite this, some species manage to persist in urban areas, as is the case with African Crowned Eagles Stephanoaetus coronatus in KwaZulu-Natal, South Africa. As relatively little is known about African Crowned Eagle nestling diet, especially about how it changes with nestling age, we investigated this with nest camera-traps. We analysed temporal changes in prey composition and biomass delivery during the nestling stage. We also recorded which adults provisioned and attended the nest. The main prey fed to nestlings were Rock Hyrax Procavia capensis and Hadeda Ibis Bostrychia hagedash. Adult males did most of the food provisioning, especially at the start of the nestling period. We found a decrease in total prey number and biomass with nestling age. This may be caused by changing requirements of nestlings. Furthermore, delivering fewer prey at later nestling stages may be a facilitating mechanism to enhance fledging of the nestling. Although the total number of prey brought to the nest decreased, we found an increase in numbers of Vervet Monkey Chlorocebus pygerythrus in the diet with nestling age. This indicated an increase in larger prey being delivered to the nests as the nestling aged. We suggest that this could be caused by increased participation in hunting by the larger female as her nest attendance time decreased as the nestling aged. We conclude with emphasising the importance of protecting the Durban Metropolitan Open Space System (D’MOSS) zones for the persistence of this Near Threatened raptor species, and populations of its prey in urban areas for its breeding success.     

Prey of White-throated Hawks (Buteo albigula) in the southern temperate forest of Argentina

ABSTRACT.   We examined the diet of White-throated Hawks ( Buteo albigula ) during the incubation and nestling periods in the southern temperate forest of Argentina. Pellets ( N = 74) and prey remains ( N = 59) were collected at 10 nests from 1998 to 2003, and preys delivered to two nests were monitored during the 2001–2002 breeding seasons. White-throated Hawks fed on small mammals, birds, lizards, and insects. The three methods of identifying prey (pellets, prey remains, and direct observation) produced different results. All types of prey except large birds were detected in pellets, and arthropods may have been over-represented in pellets due to secondary consumption. No remains of either arthropods or reptiles were identified among prey remains collected at nest sites and, during nest observations, we were unable to identify many of the prey items delivered by adults. Our results indicate that accurate determination of the diet of White-throated Hawks requires more than one method of identifying prey.     

First nest description, breeding, ranging and foraging behaviour of the Short-legged Ground-Roller Brachypteracias leptosomus in Madagascar

The secretive, endemic Short-legged Ground-Roller Brachypteracias leptosomus was studied from October 1996 to February 1997 on the Masoala Peninsula, northeast Madagascar. Several vocalizations were associated with contact, courtship feeding and food solicitation. One study pair ranged within an area of 19.1 ha and spent 90% of their time together. They used small trees for foraging and resting, and durations of perch time averaged 9.8 min. Of the 229 identified prey items recorded, 88% were invertebrates and 12% vertebrates. The first described nests for this species were observed in December 1996 and January 1997. The first nest was in a natural tree cavity 18.1 m above the ground in a 133-cm diameter-at-breast height (dbh) Weinmannia sp., and it contained at least one egg. This nest failed on 1 January 1997 when a swarm of Honey Bees Apis mellifera took over the cavity. On 7 January, the pair began excavating another nest 22 m above the ground in a 174-cm dbh Canarium madagascarense , in the root mass and decayed material of epiphytes and below a 1 -m diameter forked branch. Incubation lasted between 22–26 days and the nestling period was 30 days. One young fledged in March 1997.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

SOME NOTES ON THE CAPE GANNET,Morus capensis

Tarboton, W. R. 1978. Breeding of the Little Banded Goshawk. Ostrich 49:132-143.

The behaviour and vocalizations of a pair of Little Banded Goshawks Accipiter badius during part of their breeding cycle is described. Both sexes built the nest. Two eggs were laid three days apart. The first egg was incubated for 52% of the day, but this increased to 90% when the clutch was complete, of which the female's share was 86% and the male's 4%. The second egg hatched after 29 days, 18 h. The female did not hunt during the incubation or early nestling period and was fed by the male who brought her, on average, 7,0 food objects per day. Lizards formed 73% of the 91 identified prey objects, and small birds, 24%. The female and chick, when 16 days old, were killed by a predator on the nest at night.     

Breeding biology,diet and vocalization of the Helmet Vanga,Euryceros prevostii,on the Masoala Peninsula,Madagascar

La Morco, G. & Thorstrom, R. 2000. Breeding biology, diet and vocalization of the Helmet Vanga Euryceros prevostii on the Masoala Peninsula, Madagascar. Ostrich 71 (3&4): 400-403. The endemic Helmet Vanga, Euryceros prevostii, was studied from October to December 1997, with incidental observations from October to December 1993-1997, on the Masoala Peninsula, northeastern Madagascar. Three types of vocalizations of this species were associated with territorial, alarm and contact calls, respectively. Nests were open cups placed in the forks of trees or at the apex of small trees. Nests were composed of interlaced grasses in the nest bawl, woven plant fibres supporting the centre of the nest and lichen/moss on the exterior portion of the nest. Average clutch size was 2.7 (n = 6 nests). During incubation, either adult was on the nest 99% of the time. The reproductive output was 1.8 young fledged per nest (n = 4 nests) with 75% nest success (3/4). Of the 106 prey items recorded, 91% were invertebrates and 9% verte-brates. Cockroaches, butterflies, moths, crickets, katydids and beetles represented the most numerous insect prey taken, representing 73% of the identified prey. [A French translation of the abstract is provided on p. 403.]     

4. GAME RESERVES IN SOUTHERN AFRICA

Steyn, P. &; Grobler, J. H. 1981. Breeding biology of the Booted Eagle in South Africa. Ostrich 52:108-118.

The Booted Eagle Hieraaetus pennatus is a breeding visitor to the Cape Province of South Africa, wintering mostly in Namibia on present knowledge. Palaearctic birds probably also reach the Cape but arrive later. Two nests in different localities in the Cape were studied. The birds breed soon after arrival. Both sexes build the nest on a cliff ledge. Incubation, which lasts 40 days, is done mostly by the female. The female spends most of her time on the nest during the first four weeks of the nestling period, but considerably less time thereafter. The male provides nearly all the prey until near the end of the nestling period, and helps to feed the young. Details of nestling growth and behaviour and of parental care are given. The nestling period was 50 and 54 days in two cases. Post-nestling dependence is about two months. Prey preferences in the two study areas were very similar: 54% birds, 33% lizards and 13% rodents. Breeding biology in South Africa is basically the same as that of Palaearctic populations, with the main differences being the habitat and cliff nest site.     

Individual Consistency in Parental Effort Across Multiple Stages of Care in the House Sparrow, Passer domesticus

In many avian species in which biparental care is provided to offspring, substantial variation exists within members of each sex in the level of effort contributed to various forms of parental care. Questions remain as to whether individuals that contribute more toward one parental activity also contribute more toward other activities in which they participate. We examined the contributions of male and female house sparrows (Passer domesticus) to three forms of parental care: incubation, nestling provisioning, and nest defense, and compared the investments made by individuals at each stage of care relative to other same‐sexed parents. In both males and females, nestling feeding rates were positively associated with time spent incubating, but no relationships were found between measures of nestling feeding and nest defense. The predictability of an individual's feeding behavior based on earlier incubation efforts may make incubation a good stage for individuals to evaluate the parental abilities of their partners.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Why do Both Parents Incubate in the Kentish Plover?

Incubation by both parents is a common parental behaviour in many avian species. Biparental incubation is expected if the survival prospects of offspring are greatly raised by shared care, relative to the costs incurred by each parent. We investigated this proposition in the Kentish plover Charadrius alexandrinus, in which both parents incubate the clutch, but one parent (either the male or the female) usually deserts after hatching of the eggs. We carried out a mate‐removal and food supplementation experiment to reveal both the role of the sexes and food abundance in maintaining biparental incubation by removing either the male or the female from the nest for a short period of time. In some nests we provided supplementary food for the parent that remained at the nest to reduce the costs of incubation, whereas other nests were left unsupplemented. Although males spent more time on incubation after their mate had been removed, females’ incubation did not change. Notwithstanding the increased male incubation, total nest attentiveness was lower at uniparental nests than at biparental controls. However, incubation behaviour was not influenced by food supplementation. We conclude that offspring desertion during incubation is apparently costly in the Kentish plover, and this cost cannot be ameliorated with supplementary food.     

四川南充市区珠颈斑鸠的繁殖生态学和巢址选择

2002年11月~2004年4月在四川省南充市区内对珠颈斑鸠(Streptopelia chinensis)繁殖生态和巢址选择进行了研究。结果表明:珠颈斑鸠3月初开始求偶交配,求偶行为复杂,有“婚飞”行为;雌雄参与筑巢,营巢期7~8 d。影响巢址选择的主要因素有6种:栖位与巢周隐蔽因子、巢下隐蔽因子、光照因子、人为活动因子、食物因子和营巢树因子;窝卵数2枚,雌雄轮流孵卵,孵卵期17~18 d,孵化率86.67%;雌雄均参与育雏,育雏期18~20 d,雏离巢率73.08%,繁殖生产力1.82,种群育雏高峰期为7月和8月中上旬。     

How does an increase in minimum daily temperatures during incubation influence reproduction in the great tit Parus major?

Temperature variation affects all life stages of organisms, especially early development, and considering global warming, it is urgent to understand precisely its consequences. In egg‐laying species, incubation behaviour can buffer embryo developmental temperature variation and influence offspring development. We experimentally investigated the effect of an increase in minimum daily nest temperature during incubation in the great tit Parus major, by placing a hand warming pad under the nest in the evenings. As compared to controls, the experimental treatment increased nest temperature at night by an average of 4°C, and this increase carried over to the following day. We measured the consequences of this mainly nocturnal temperature increase during incubation on 1) parental behaviour (incubation and nestling feeding), 2) parental health (quantified by body condition, immune status, physiological and oxidative stress) and 3) reproductive success (nestling body condition, growth, i.e. mass gain, hatching and fledging success, and nestling immune status, physiological and oxidative stress). This study yielded three major results. First, we found that heating the nest did not change the duration of incubation as compared to controls. Second, increasing nest temperature during incubation decreased nestling feeding behaviour but did not affect parental health in terms of body condition, immune status, physiological and oxidative stress. Third, nestling mass at hatching was greater but nestling mass gain was slower in heated nests than in control nests, resulting in similar fledging mass. The present study demonstrates that increased environmental temperatures during incubation influenced nestling development in the great tit and especially hatchling mass, which might produce long‐term life history consequences.     

Is Mate Provisioning Predicted by Ornamentation? A Test with Northern Cardinals (Cardinalis cardinalis)

Male birds of many species feed their mates during courtship and incubation. The amount of food provided can be substantial and even essential for successful reproduction in some species, and can influence female nest attentiveness in many others. Additionally, mate provisioning may predict later nestling feeding rates. Females may thus benefit from being able to determine male provisioning effort. We assessed the expression of several ornaments, known to indicate condition in male northern cardinals (Cardinalis cardinalis), and compared these with mate provisioning rates, nestling feeding rates, and nest attentiveness. We found that male ornamentation may not be indicative of mate provisioning rates. Mate provisioning rate did not co‐vary with reproductive success, male feedings to nestlings, or nest attentiveness of females. However, females which were fed more often during incubation tended to provision nestlings less. Reduced female parental effort following extensive incubation feeding may be indicative of females using incubation feeding to assess future male parental effort. Male hormonal condition that favors high rates of nestling provisioning may be a proximate cause of mate provisioning during incubation, even in the absence of selection, favoring high rates of mate provisioning. Both sexes may have capitalized on this unselected behavior.     

Predation risk of eggs and nestlings relative to nest‐site characteristics of the Bull‐headed Shrike Lanius bucephalus

Appropriate nest‐site selection is one of the most important ways to minimize loss of reproductive investment due to predation. We determined the environmental characteristics associated with nest predation during the incubation and nestling periods of arboreal nesting Bull‐headed Shrikes on the oceanic Minami‐Daito Island where the predator community has low species diversity and includes only three introduced mammals: Ship Rat Rattus rattus, Japanese Weasel Mustela itatsi and Feral Cat Felis catus. Egg predation declined with increasing grassland cover around nests, whereas nestling predation declined with increasing nest concealment and nest height. Our results suggest that effective nest‐site characteristics for avoiding nest predation differ during the incubation and nestling periods and are dependent on the predator species and their search strategies, at least in habitats with low predator species diversity.     

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.     

Foraging behaviour of nest predators at open-cup nests of woodland passerines

Nest predation patterns and processes cannot be understood without studying the behaviour of predators. I videotaped the behaviour of 22 species of predators at 171 depredated nests of 13 passerine species, in woodland in the Czech Republic. About 32% (60/187) of all events occurred during the night; mammals accounted for 95% (57/60) and 22% (28/127) of nocturnal and diurnal predation, respectively. About 67% (57/85) of mammalian predation, but only 3% (3/102) of avian predation, occurred during night. Multiple predations by the same species were detected in at least 7% (6/82) and 42% (37/88) of nests depredated by mammals and birds, respectively. Martens Martes martes/foina took nest content mostly all at once; birds (mainly Jay Garrulus glandarius) revisited partially depredated nest during 1–4 consecutive days. Martens stayed at the depredated nest about five times longer than Jays. Martens spent similar time at nests with eggs and nestling, while Jays stayed about twice longer at nests with eggs. Mammals consumed eggs always at the nest (23/23), but took nestlings away in at least 48% (31/64) cases. Birds took the eggs and nestling away in at least 31% (18/58) and 76% (71/94) cases, respectively. Predator visits to active nests without taking the content, repeated partial predation and revisitation of previously depredated nests suggest an effect of memory on predator’s foraging behaviour.