Size-biased allocation of prey from male to offspring via female: family conflicts, prey selection, and evolution of sexual size dimorphism in raptors

In birds with bi-parental care, the provisioning link between prey capture and delivery to dependent offspring is regarded as often symmetric between the mates. However, in raptors, the larger female usually broods and feeds the nestlings, while the smaller male provides food for the family, assisted by the female in the latter part of the nestling period, if at all. Prey items are relatively large and often impossible for nestlings to handle without extended maternal assistance. We video-recorded prey delivery and handling in nests of a raptor with a wide diet, the Eurasian kestrel Falco tinnunculus, and simultaneously observed prey transfer from male to female outside the nest. The male selectively allocated larger items, in particular birds and larger mammals, to the female for further processing and feeding of nestlings, and smaller items, in particular lizards and smaller mammals, directly to the nestlings for unassisted feeding. Hence, from the video, the female appeared to have captured larger prey than the male, while in reality no difference existed. The female’s size-biased interception of the male’s prey provisioning line would maximize the male’s foraging time, and maximize the female’s control of the allocation of food between her own need and that of the offspring. The male would maximize his control of food allocation by capturing smaller prey. This conflict would select for larger dominant females and smaller energy-efficient males, and induce stronger selection the longer the female depends on the male for self-feeding, as a proportion of the offspring dependence period.     

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.     

The relative influence of prey abundance and co-breeders on the reproductive performance of polyandrous Pale Chanting-goshawks

This study investigates if the reproductive performance of polyandrous Pale Chanting-goshawks, Melierax canorus, is governed by the abundance of dominant rodent-prey species or a co-breeding male participating fully in prey being delivered to the female and young. Polyandrous trios in prey-rich habitat, the only habitat where these trios occurred, failed to produce more offspring than monogamous pairs, but attempted more second broods in years of high prey abundance. The higher the prey-delivering rates at the nesting sites by either one monogamous male or two polyandrous males (combined effort), the higher the proportion of time their females spent in close proximity to the nest. Since there were no significant differences between the body masses of nestlings fledged by polyandrous and monogamous groups, higher delivery rates relieved females from away-from-the-nest hunting duties and permitted them to feed on prey not fed to offspring. Although breeding in polyandrous trios held fitness benefits for females, e.g. to accumulate the necessary body reserves for laying a second clutch in years of high prey abundance, it is still not clear how males benefited. The reproductive performance of each group was strongly linked to the abundance of the dominant, otomyine rodent prey (Otomys unisulcatus and Parotomys brantsii) and it was therefore prey abundance, and not the number of males delivering prey to the female and nestlings at each nesting site, that controlled the reproductive performance of cooperative-breeding Pale Chanting-goshawks.     

First nest descriptions,nesting biology and food habits for Bernier's Vanga,Oriolia bernieri,in Madagascar

Four nests of the rare and endemic Bernier's Vanga, Oriolia bernieri, were discovered; one in 1997, one in 1998, and two in 1999 on the Masoala Peninsula, northeastern Madagascar. At the 1998 nest, the female made 189 visits with 186 deliveries of nesting material during 34.6 h of observation. The female spent 9.2% (194.2 min) of the observation time building the nest while an immature male delivered nest material six times and spent 3.2 min at the nest placing the material. Nesting material included: 67.2% (125) decomposed root material, 24.7% (46) palm fibres, 6.5% (12) dry leaves, 1.1% (2) moss, and 0.5% (1) white plant dawn. In 41.0 h of observation during the incubation period the female incubated for 53% (21.7 h) of the time, the adult male for 32.3% (13.2 h), the immature male for 4.3% (1.8 h), and the nest was unattended for 10.4% (4.3 h). This breeding attempt foiled on day 13 of incubation when a Madagascar Harrier-Hawk, Polyboroides radiatus, ate the egg(s). At one of the 1999 nests, the incubation and nestling periods were 17 days each. Three young fledged during the middle of November. Of the 82 identified prey items recorded during the nestling period, 91% were invertebrates and 9% vertebrates. Spiders, crickets, cockroaches, and geckos represented the most numerous prey taken, totaling 77% of the identified prey.     

Diversionary feeding and nestling diet of Hen Harriers Circus cyaneus

Capsule: Diversionary feeding reduced Hen Harrier Circus cyaneus nestlings’ natural food intake by half. Red Grouse Lagopus lagopus scotica chicks constituted 0–4% of all nestling food items. Annually, this reduced annual grouse chick production by 0–6%.

Aim: To quantify proportions of diversionary and natural food (including grouse) delivered to Hen Harrier nestlings in relation to brood size, male status and natural prey abundance.

Methods: We recorded diversionary food provisioned to 25 Hen Harrier broods (2008–15) and studied the diet of 15 broods using observations from hides, nest cameras and regurgitated pellet analysis. Variation in nestling diet was analysed using compositional analysis.

Results: Hen Harriers took 76% of diversionary food provided. Depending on assessment method, average nestling diet was 44–53% diversionary food, 39–55% natural prey (including 24–45% passerines, 4–15% small mammals, 0–4% grouse chicks) and 0–9% unknown items. The amount of diversionary food consumed was not influenced by male status, brood size or natural prey abundance. The number of Red Grouse chicks delivered annually was 34–100% lower than expected under unfed conditions, however, the confidence intervals associated with these estimates were large.

Conclusion: Diversionary food influenced Hen Harrier nestling diet and reduced the number of Red Grouse chicks taken relative to modelled predictions. It may help reduce conflict between Hen Harrier conservation and Red Grouse shooting, but only if overall grouse productivity is thereby maintained or increased.     

Feeding ecology and specialist diet of critically endangered Ridgway's Hawks

Ridgway's Hawks (Buteo ridgwayi) are critically endangered forest raptors endemic to the Caribbean island of Hispaniola, with ～100 pairs remaining in the world. The species is ecologically little known yet such studies are important for understanding critical habitat needs and population dynamics. We studied the provisioning behavior of adults at 22 nests on the northeast coast of the Dominican Republic from 2005 to 2008. Mean brood size was 1.80 ± 0.45, and the mean number of fledglings per nest was 1.10 ± 0.97. We found that 80% of the prey items delivered to nestlings were reptiles, with lizards accounting for 65% of the prey and those in the genus Celestus accounting for nearly 35% of prey. Other prey items included snakes (14%), rats (9%), and smaller proportions of birds, frogs, and centipedes. The number of prey items and amount of biomass delivered to nestlings did not vary with brood size, but adults delivered more prey to 3‐ to 5‐week‐old nestlings and more biomass to 5‐week‐old nestlings. Food delivery rates did not differ between successful or failed nests, suggesting that food availability did not influence nest outcome. Given that most prey items delivered to nestlings in our study were reptiles, conservation strategies developed for Ridgway's Hawks (e.g., translocations and habitat conservation) should take into consideration their specialist reptile diet.     

Video nest monitoring reveals male coloration-dependant nest predation and sex differences in prey size delivery in a bird under high sexual selection

Increased predation risk should select for reduced parental activity to decrease the probability of visually hunting predators discovering the nest. Parental activity and conspicuousness are known to increase predation risk. Here, we test for sex differences in parental visitation rate (number of visits), time seen at the nest (time at the nest × adult visibility), and food delivery (prey size) using continuous video recordings at nests. We test the role of these variables for predation outcome in the Superb Fairy-wren (Malurus cyaneus). The study species is sexually dimorphic: males have iridescent blue plumage while females have brown plumage. The results showed that nest predation was predicted by male time seen at the nest (but not visitation rate), but not female time seen at the nest (or visitation rate). Contrary to our expectation that males would have lower visitation rates than females, our analysis of video images showed that male and female visitation was comparable but that males consistently brought smaller prey items to the nest than females. These findings are discussed in the light of morphological differences between male and female beak size. We conclude that sexual selection has favoured conspicuous male signalling in this system, and that natural selection should select for reduced parental care for the conspicuous sex.     

NOTES ON THE CAPRIMULGIDAE OF THE KAFUE BASIN

During 2002–2007, we assessed the status of the Purple Heron Ardea purpurea in Numidia, Algeria by surveying all the major wetlands in the region. We located six distinct breeding sites; four of these were not previously known. We also investigated nest site selection and determined the species’ reproductive success at Dakhla, a dunary pond during two successive years (2006 and 2007). The egg laying period was comparable to that reported for southern Europe (March-May) and a seasonal change of breeding success was recorded. A growth curve was derived for developing nestlings and the impact of an ectoparasite (mite) infestation upon nestling's growth was considered. We also examined 73 food boluses regurgitated by nestlings and identified 329 prey items. Fish, mainly Gambusia holbrookii and Cyprinus carpio, dominated by mass, whereas insects, mainly aquatic Coleopteran larvae and aeshnids (Odonata), were the most frequent prey (67.1%). Loss of habitat is identified as a major threat to the future of colonial herons in Algeria.     

Parental differences in brood provisioning by Hen Harriers Circus cyaneus

Capsule Females varied their provisioning patterns according to brood age and brood size, whereas males did not.

Aims To quantify how parents balance the needs of their offspring for food and protection.

Methods We studied 13 nests from hides and spent on average 101 hours per nest monitoring prey types, provisioning rate and the time spent at the nest by both sexes in relation to brood size and brood age.

Results Males always provided more food than females. Males brought similar amounts of prey items irrespective of brood size and nestling age, whereas females brought more prey and bigger items to larger and older broods. Females spent less time brooding larger broods, particularly early on.

Conclusions Hen Harrier parents share the provisioning burden, with each parent delivering prey as a function of brood care requirements, hunting capability and the behaviour of the other parent.     

Evidence of prey depletion around lesser kestrel Falco naumanni colonies and its short term negative consequences

Predation may reduce prey numbers in such extent that prey may be depleted, which has negative effects on predator populations. Prey depletion would be more likely when the number of predators increase and/or concentrate their activity in a certain area, as is the case of colonial birds. As a matter of fact, food depletion due to intraspecific competition is considered a major cost of coloniality, and several studies have shown indirect evidence of this. However, no direct measures of food depletion have been provided along with its consequences for the fitness of the colony inhabitants. We carried out a field study with the lesser kestrel Falco naumanni, a raptor that breeds in colonies ranging from two to dozens of pairs. During the nestling period we sampled the main prey of the kestrels around different sized colonies at increasing distances. At the same time, we recorded hunting distances and prey delivery rates to the nest. In addition, we monitored the reproductive success in colonies of different sizes. Lesser kestrels feed their nestlings mainly with grasshoppers and these prey became depleted through the season in the surroundings of the large colonies. Prey depletion made kestrels fly longer distances to forage and prey delivery rates to the nest decreased. Lower feeding rates were not compensated by bringing larger prey, hence, the net amount of energy provided to the chicks decreased with the date in large colonies. By contrast, none of this occurred around small colonies, where both prey abundance and hunting distance remained constant throughout the season. As a consequence, the seasonal decline in the reproductive success (number of fledglings and fledgling body condition) was greater the larger the colony. Thus, these results evidence that food depletion and its fitness costs are related to colony size, as they are suffered by the kestrels breeding in large colonies but not by those settled at small ones. Finally, the consequences of prey depletion on the demographic dynamics and the regulation of colony size are discussed.     

Fitness consequences of male provisioning of incubating females in a desert passerine bird

Male provisioning of incubating females can increase reproductive success by maintaining physiological condition of females and consistency of incubation. The effects of male provisioning on the maintenance of incubation temperature and embryo development should be particularly pronounced in environments where ambient temperature exceeds the tolerance of unincubated eggs and where consistency of female incubation might be particularly important for hatching success. Here, we investigated the reproductive consequences of incubation feeding in a desert population of House Finches (Carpodacus mexicanus) in southwestern Arizona. We found that greater nest attentiveness by females was related to higher minimum incubation nest temperature, that in turn was closely associated with hatching success. Only 44% of males regularly provisioned their incubating females. Although provisioned females maintained higher incubation temperature and took fewer incubation breaks than non-provisioned females, overall, male provisioning did not influence incubation dynamics or hatching success. Further, a male’s incubation feeding rate did not correlate with male provisioning of nestlings. These results corroborate the finding that, in male House Finches, neither provisioning of incubating females nor pre-incubation courtship feeding are associated with increases in circulating pituitary prolactin––the hormone regulating male provisioning of nestlings. We suggest that incubation provisioning by male might be a component of pair maintenance behavior and that variation in male incubation behavior is best understood in relation to asymmetries in residual reproductive values between the mates.     

Site fidelity and home range variation during the breeding season of peregrine falcons (Falco peregrinus) in Yamal,Russia

Many different ecological factors affecting the size, use, and spatial configuration of home ranges have been investigated, yet the chronology of the breeding cycle has been relatively under studied. Here, we studied peregrine falcons (Falco peregrinus) to describe variation in home range within and between breeding seasons in the Yamal peninsula, a region of the Russian Arctic with a high breeding density of peregrines. We used satellite telemetry to investigate variation in home range at different stages of the breeding cycle during three breeding seasons (2009–2011). We fitted Argos satellite transmitters to 10 breeding peregrines (nine females and one male) and two male fledglings. All breeding females showed fidelity to the region of the southwestern Yamal peninsula, but they were not necessarily faithful to their specific breeding ranges with 33 % dispersing to new ranges up to 40 km away. The population of peregrines in our study area was relatively synchronous in their breeding chronology, with clutches initiated in close synchrony in early June despite the birds arriving on their breeding ranges ca. 3 weeks earlier. The average home range size for breeding females was 98 km² (95 % Maximum Convex Polygon). Over the breeding season, the home range area utilized by females increased in the late nestling period and again after the chicks fledged. Expansion of the home range coincided with changes in behavior associated with parental care, resulting in greater activity and more time spent away from the nest area when the female began hunting to provision nestlings and fledglings.     

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.     

NOTICES

Steyn, P. 1975. Observations on the African Hawk-Eagle. Ostrich 46:87-105.

Observations extending over 12 years were made on two pairs of African Hawk-Eagles Hieraaetus spilogaster at Essexvale, Rhodesia. Details on various aspects of adult behaviour are given, particularly on hunting methods and calls. Nest repair usually took about 4–5 weeks. and limited observations indicated that the male does most of the work. Incubation is done mostly by the female, the male relieving her when he brings prey. The incubation period is 43 ± 1 day. Details are given of parental behaviour during the fedging period; time on the nest showed a progressive decline although both adults still perched in the. nest tree a great deal. The male provided most of the prey. The growth and behaviour of the eaglet Is described; usually the eaglet becomes a “brancher” before its first flight which, in four cases, occurred between 61–71 days. Post-of edping attachment to the nest lasted about three weeks in one case. Frief mention is made on the development of adult plumage. Birds made up 74%. mammals 25% and reptiles 1% of 104 prey items recorded at Essexvale. Gamebirds are preferred. The two nests had completely different breeding histories. One pair reared no young while the replacement rate of the other pair was 0,83 voung/pair/year. The combined replacement rate of the two pairs was 0,48 young/pair/year. There appear to be no records of two eaglets being reared together in southern Africa.     

Nesting behavior of the semi-aquatic spider wasp,Anoplius eous,which transports its prey on the surface film of water (Hymenoptera,Pompilidae)

Anoplius eous Yasumatsu (Hymenoptera, Pompilidae) exhibits some outstanding nesting behavior. Females excavate a unicellular or multicellular nest in wet ground, or dig a single-celled nest in rotten wood, or build clustered mud cells in narrow spaces between walls of such substances as wood or vinyl sheets. The latter nest-type has been unknown within the subfamily Pompilinae. Females prepare a nest-cell before hunting, and construct it further after hunting, leaving the prey near the nest (behavior-formula: IVPTIOC). They transport their prey backward on the ground, grasping it in their mandibles by any part of the legs, and forward on the surface film of water or on the ground, grasping it by the middle part of the 1st or 2nd legs. Their only prey is the adult female semi-aquatic spider,Pardosa pseudoannulata (Lycosidae). These nesting and provisioning behavior patterns are compared with those of other pompilids. The position of the present species in the behavioral evolution of the Pompilidae is suggested.     

Brutzeitliche Beutewahl beim Seggenrohrs?ngerAcrocephalus paludicola

Summary Aquatic Warblers inhabitCarex Marshes (Magnocaricia) exceptionally rich in arthropods. Contrasting with the otherAcrocephalus species their mating system is some form of polygyny resp. promiscuity and the female alone feeds her nestlings. At 17 nests females fed large prey items compared with the arthropod fauna of the vegetation. Food was collected close to the nest and feeding rate proved to be high. Therefore richness of arthropods in the vegetation seems to form a prerequisite to this mating system untypical amongAcrocephalus warblers.     

Daily patterns of nest visits are correlated with ambient temperature in the Northern Wheatear

The diurnal pattern of foraging behaviour in birds is commonly linked to the risk of starvation and predation. During the breeding season, when offspring place additional demands on the quantity of food adults need to collect, diurnal foraging patterns may be influenced to a greater extent by food availability because adults need to balance starvation–predation risks not only for themselves but also their offspring. We used data-loggers to measure diurnal variation in rates of nest visits to 7- to 10-day-old nestlings of the insectivorous Northern Wheatear Oenanthe oenanthe in Swedish farmland habitat. Adults provisioned the nestlings with food for an average 19.5 h each day (n = 11 pairs), with visitation rates sharply increasing during the morning, remaining relatively stable between 0800 and 1700 hours, and then declining in the evening. This pattern is similar to daily temperature fluctuations and is consistent with the hypothesis that nest visit rates are a function of temperature-dependent prey availability. Higher rates of nest visits on warm days support this correlation between temperature and offspring feeding. However, without data on the rate of self-feeding by adults it is difficult to distinguish between patterns of nest visits being driven by food availability, or starvation–predation constraints being imposed on the parents; thus, future studies of patterns of offspring feeding should collect data on the self-feeding behaviour of the adults.     

Male Rock Sparrow (Petronia petronia) Nest Defence Correlates with Female Ornament Size

We investigated the relationship between male nest defence and female breast patch size in an alpine population of rock sparrow (Petronia petronia) in northern Italy. We presented a mounted weasel (Mustela nivalis), a common nest predator, to 28 pairs breeding in nest boxes, with 12–13‐d‐old nestlings, and measured the intensity of male and female defence reaction. We measured the frequency of attack flights, intensity of alarm calling and total time spent in view, and then combined these for each individual, in a single defence factor by principal component analysis. All the females arrived to defend the nest while only 21 males arrived, and females defended the nest more intensely than males. We analysed, by stepwise regression, the relationship of male defence factor to female behaviour and phenotype (breast patch size, a measure of quality) and brood properties (size, mass, phenology). Male defence factor was significantly related only to female breast patch size. We argue that male rock sparrows apparently make parental investment decisions according to their mate's quality, and examine possible alternative hypotheses.     

Patterns of nest provisioning and parental investment in the solitary digger wasp Ammophila sabulosa

Abstract.

• 1 In nest-building wasps there is a continuum from species that provision each offspring cell with one large prey item (single provisioning) to species that provision each cell with many small items (multiple provisioning). The significance of number of prey per cell was examined during two seasons in Ammophila sabulosa (L.), a species that provisions between one and five caterpillars per cell.
• 2 There was no difference between the total weight of prey in singly-provisioned and multiply-provisioned cells: individual caterpillars placed in multiply-provisioned cells were smaller.
• 3 Small caterpillars were captured and transported to the nest faster than large ones.
• 4 If the first prey provisioned was large, the cell was usually then permanently closed. If the first prey was small, additional prey were captured, and females appeared to become more selective: additional prey were rarely large.
• 5 It is suggested that the flexible provisioning strategy used by A.sabulosa allows a wide range of prey sizes to be utilized without affecting maternal control of offspring size and sex. This could be important when available prey sizes vary temporally and spatially.
• 6 There was a male-biased first generation investment ratio. Female-producing cells were provisioned with a higher total prey weight than male-producing cells, and conversion of prey weight to adult weight was more efficient for females.
• 7 Total prey weight provisioned is probably a good indirect measure of parental time investment in A.sabulosa. Other measures, particularly number of prey provisioned, will be less accurate.

     

The use of agricultural, open and forest habitats by juvenile Mauritius Kestrels Falco punctatus

Knowledge of tropical raptor habitat use is limited and yet a thorough understanding is vital when trying to conserve endangered species. We used a well studied, reintroduced population of the vulnerable Mauritius Kestrel Falco punctatus to investigate habitat preferences in a modified landscape. We constructed a high resolution digital habitat map and radiotracked 13 juvenile Kestrels to quantify habitat preferences. We distinguished seven habitat types in our study area and tracked Kestrels from 71 to 130 days old during which they dispersed from their natal territory and settled within a home‐range after reaching independence. Mean home‐range size was 0.95 km² characterized by a bimodal pattern of intensity around the natal site and post‐independence home‐range. Compositional analysis showed that home‐ranges were located non‐randomly with respect to habitat but there was no evidence to suggest differential use of habitats within home‐ranges. Native and semi‐invaded forest and grassland were consistently preferred, whereas agriculture was used significantly less than other habitats. No difference was found between the available length of edge dividing native forest and grassland within a home‐range when compared to that available within a 2.35‐km buffer around their nest‐site, based on the maximum distance a juvenile was found to disperse. Repeating the analysis in three dimensions gave very similar results. Our results suggest that Mauritius Kestrels are not obligate forest dwellers as was once thought but can also exploit open habitats such as grassland. Kestrels may be using isolated mature trees within grassland as vantage points for hunting in the same way as they use the natural stratified forest structure. We suggest that the avoidance of agriculture is partly due to a lack of such vantage points. The conservation importance of forest degradation and agricultural encroachment is highlighted and comparisons with the habitat preferences of other tropical falcons are discussed.