4. ONE YEAR'S CENSUS AND STUDY OF BIRDS IN 2½ ACRES OF ALEANY BUSHVELD. (Part 2.) April, 1945, to March, 1946

Hooded Vultures Necrosyrtes monachus are critically endangered but little is known of their year-round use of nests or whether other species usurp Hooded Vulture nest sites. We investigated visitation rates by Hooded Vultures and other species (including potential nest predators and usurpers) to examine their effect on Hooded Vulture breeding success. We present observations of 33 species recorded by camera traps at 12 Hooded Vulture nests over a total of 93 nest-months (2 095 nest-days). Several pairs of Hooded Vultures visited their nests regularly during the non-breeding season, some adding nesting material, highlighting that pairs visited their nest(s) year round. Egyptian Geese Alopochen aegyptiaca, potential usurpers of raptor nests, were present at occupied and unoccupied Hooded Vulture nests, but we recorded no usurpation of nests by Egyptian Geese and they had no impact on vulture breeding success. Hooded Vulture breeding failure was linked to two species only: camera-trap imagery recorded one case of predation of a vulture egg by a Chacma Baboon Papio ursinus, and one case of a Martial Eagle Polemaetus bellicosus predating a vulture nestling. We recommend expanding the Hooded Vulture nest monitoring programme to include more pairs.     

Anti-social and social behaviour of adolescent yellow-rumped caciques (Icterinae: Cacicus cela)

In an individually marked population in southeastern Peru, males and females of the polygynous, colonial yellow-rumped cacique delayed successful breeding until they were 3 years old. Adolescent (2-year-old) females competed for nest sites, built nests, laid fertile eggs, incubated, and sometimes fed nestlings, but always abandoned nests before they fledged young. Females that completed nests during their adolescent years were also more successful in establishing nest sites the following year. Adolescent females that failed to establish nest sites and those that lost nests to predators tended to leave the study area altogether. By attempting to nest early, adolescent females may facilitate future nesting attempts by forming coalitions with other females and assessing nest site quality in an area. Adolescent males, on the other hand, avoided aggressive interactions with other males, but were very aggressive towards adult females and fledglings. Adolescent males attacked females in nests, chased females to and from the colony, pecked and mounted recent fledglings, and, rarely, pulled nestlings out of nests. This harassment resulted in the death of nestlings and fledglings and was energetically costly to females, which often fought adolescent males while defending their young. Adolescent males preferentially harassed nestling-feeding females, avoided egg-laying females and were never observed attempting to mate with females. Adult males rarely interfered during any of these interactions. Such anti-social behaviour may function as a way of practising adult behaviour, as a means of eliminating future competitors, or may be an epiphenomenon of rising hormone levels and therefore of no adaptive significance.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.     

The presence of kleptoparasitic fledglings is associated with a reduced breeding success in the host family in the barn owl

Fledgling birds sometimes abandon their own nest and move to neighboring nests where they are fed by host parents. This behaviour, referred to as ‘nest‐switching’, is well known in precocial birds that are mobile soon after hatching and can easily reach foster nests. In contrast, due to the difficulty of observing nest‐switching in territorial altricial birds, the causes and consequences of moving to others’ nests are poorly known in this group of birds. Nest‐switchers can be adopted by the foster parents or they can steal food from the host parents meant for their offspring, a form of kleptoparasitism, which may result in reduced breeding success of the host nest. In Israel, 12 barn owl fledglings left their natal nests and were found in 9 host nests out of 111 monitored nests (8.1%). Nest‐switchers that fledged earlier in the breeding season flew shorter distances to reach host nests probably because the density of nests with younger nestlings is higher early in the season. The number of host nestlings fledged and the percentage of nestlings fledged was lower in host nests than in nests without switchers. The occasional nest‐switchers were always older than host nestlings (respectively 80 and 50 days of age, on average) and host parents fledged fewer young when nest‐switchers occupied host nests with younger nestlings. This suggests that nest‐switchers are kleptoparasites because the presence of the older alien fledglings is associated with a lower breeding success of the host parents.     

Territorial behavior of the Korean wood-feeding cockroach, <Emphasis Type="Italic">Cryptocercus kyebangensis</Emphasis>

We investigated the territorial behavior of Cryptocercus kyebangensis from the viewpoint of offspring protection. Aggressive behavior against adult intruders occurred more frequently in resident adults with their young than those without their young. Resident adults without offspring more frequently attacked intruders of the same sex than those of the opposite sex, whereas those with offspring attacked regardless of the sex of adult intruders. In paired residents, cooperative territorial behavior occurred in a higher proportion in pairs with offspring than those without offspring. Responses of residents to nymphal intruders depended on the head capsule size of the intruders. Resident adults rarely attacked intruders smaller than their own offspring but frequently attacked larger intruders. Resident nymphs frequently attacked intruders smaller than themselves, whereas they escaped or showed jerking behavior when they encountered larger intruders. Electronic Publication     

Adult aggression during the post-guard phase in the chinstrap penguin Pygoscelis antarctica

We report an observational study of aggressive behaviour by adults during the post-guard phase in the chinstrap penguin. The study was carried out in one subcolony where all the individuals (breeders, chicks and failed breeders) were banded, as well as in nine other subcolonies where individuals were not identifiable. Breeding adults were more aggressive towards unrelated chicks when they were feeding their own chicks than in other contexts. Chicks were also attacked at high rates by adults that did not belong to their own subcolonies. The evidence suggests that some aggressions to chicks by adults may function to avoid interference from unrelated chicks during food transfers, but our data also show that aggressive interactions are common in contexts other than chick feeding. However, aggression by adults did not show any clear spatial directionality or purpose of shepherding the chicks, and these tended not to move in any particular direction when attacked.     

The effect of migration strategy and food availability on White Stork Ciconia ciconia breeding success

In the mid 1970s, the breeding populations of the migrant White Stork Ciconia ciconia were close to extinction in the northeastern region of France (Alsace). A re-introduction project was implemented, resulting in the year-round settlement of some individuals in the region, which rely on additional food supplied by humans during the winter. Today, both resident and migrant birds breed in the same areas and take food from rubbish dumps and humans (farmers). The effects of these anthropogenic influences, altering Stork behaviour, on Stork reproductive success are not known. The aim of this study was to test the influence of bird status (resident vs. migrant) and food availability (control nests vs. nests that benefit from high food supply) on reproductive success. In control nests, the mean laying date was earlier in resident than in migrant White Storks. There was also a clear seasonal decline in clutch size. For all nests, the numbers of eggs and hatchlings were higher in resident birds than in migrants, which can be attributed to the earlier breeding of resident Storks. The large broods of resident birds showed a high mortality rate, leading to the same fledgling success (fledglings/hatchlings) and number of fledglings as in migrants. Fledgling success and the number of fledglings were higher for nests close to a reliable food supply. In summary, although resident birds can breed earlier and produce more eggs than migrants, we found no advantage in terms of number of fledglings. The higher mortality rate of chicks found in pairs with a large brood could be caused by the deterioration of their habitat. Thus, the year-round settlement of Storks may not present a biological advantage if the quality of their habitat is not guaranteed by the conservation of their grasslands.     

Parental investment asymmetries of a globally endangered scavenger: unravelling the role of gender,weather conditions and stage of the nesting cycle

ABSTRACT

Capsule: In Egyptian Vultures Neophron percnopterus, both sexes invested similar parental effort throughout the breeding period. However, there was variation in the degree of intensity of parental care during some stages of the breeding period, suggesting that sex-role specialization exists for some activities.

Aims: To quantify parental care behaviour of Egyptian Vultures for the first time and to examine the role of sex, weather conditions, and stage of nesting cycle on breeding ecology.

Methods: We monitored 15 nests of Egyptian Vultures to analyse parental care investment. We collected data on nine different behavioural parameters/activities per sex, which were recorded throughout the entire breeding period. Variation in parental investment was analysed using generalized linear mixed models.

Results: Females invested more effort in incubation/brooding (61.45% for females and 31.54% for males) and egg turning (0.45?events/h for females and 0.37?events/h for males) while males contributed more to nest material delivery to the nest (0.67?deliveries/h for males and 0.14?deliveries/h for females). Conversely, both sexes invested the same effort in nestling attendance (21.89% for females and 21.21% for males) and food provisioning (0.28?items/h for females and 0.25?items/h for males). Furthermore, parental investment was not affected by weather, especially during critical moments such as incubation/brooding, however, changeover rate was positively related to temperature.

Conclusion: Our results suggest that, in the Egyptian Vulture, one sex is not entirely responsible for a particular task and the compensatory effort of the other mate is required. Finally, our findings indicate that major events such as incubation onset and hatching caused important shifts in the patterns of parental investment.     

Family break-up in Black and Red Kites Milvus migrans and M. milvus: is time of independence an offspring decision?

I observed the natural process of family break-up in 13 families of Black Kites Milvus migrans and five families of Red Kites M. milvus in which fledglings had been individually marked. In other broods, I performed experiments which modified the parental investment fledglings received by supplementing nests of both species with food or transferring Black Kite chicks to nests with younger or older chicks of the same species.
The time of family break-up in the Black Kite is mainly an offspring decision which is not affected by an artificial increase of parental investment. The duration of the post-fledging period was not increased in Black Kites that were given supplementary food. Chicks transferred to nests with a younger chick did not extend the post-fledging period, nor did chicks transferred to nests with an older chick shorten the post-fledging period. In Red Kites, parental investment seemed to have more influence on the timing of the family break-up. Red Kite adults invested less as parents than did Black Kites during the post-fledging dependence period, and in nests where supplementary food was given, Red Kite fledglings stayed attached for a longer period.     

The function of aggressive chases by breeding Black and Red Kites Milvus migrans and M. milvus during the post-fledging dependence period

Black and Red Kites Milvus migrans and M. milvus chase other raptors approaching their nests. The study of this behaviour during the post-fledging period suggested that it reflects mainly, but not only, anti-predator behaviour. The frequency of vigilance and aggressive chases decreased through the post-fledging period as predicted by theoretical models of nest defence. Although predation risks were similar, Black Kites invested more time chasing intruders than did Red Kites. Black Kites, unlike Red Kites, chased away intruding juveniles, which may be interpreted as a behaviour to avoid investment in unrelated fledglings. Black Kites usually nest in loose colonies where the risk of, and selection pressures against, accidental investment in unrelated fledglings is likely to be greater than for Red Kites. Differences in aggressive chases by Black and Red Kites are better related to this than to different predation risks.     

Intruders in Nests of the Spotless Starling: Prospecting for Public Information or for Immediate Nesting Resources?

The prospective behaviour for nests by conspecific intruders may be a strategy to gather public information for future reproduction or to secure resources for immediate reproduction. Although the relationship between the sex, age and breeding experience of intruders and the sex and age of residents may be crucial for understanding the significance of nest‐prospecting behaviour, a precise determination of these traits has rarely been addressed in field studies. In a free‐ranging population of spotless starlings ( Sturnus unicolor), inexperienced birds were the predominant intruders. The ratio of male‐to‐female intruders was higher among birds without previous breeding experience, although our results did not allow us to determine whether more inexperienced males or females intruded nests. The average annual rate of intrusion of either sex was strongly correlated with the number of fledglings produced in the study colony the previous year, which seems to indicate that the inexperienced intruders were, in general, yearlings. The proportion of the sexes among the intruders in each study year was correlated with the average age of male owners but not with the age of female owners. This result suggests that the presence of intruding males in the nests was affected by an increased opportunity to find vacancies in nesting territories as resident males became older. This finding supports the hypothesis that, in most cases, birds visited nests to obtain personal information about nesting resources. However, the intrusion rates were higher when there were nestlings than during the incubation period, which suggests that collecting public information was also involved in the intruding behaviour. Both sexes regularly attacked intruders, which implies that the intruders inflicted some kind of cost on the owners. Female floaters were more frequently chased, probably because they regularly lay parasitically in conspecific nests.     

On the present situation of vultures in Bulgaria: A review

Abstract

A review of the literature indicated that about 100 pairs of Egyptian Vultures, less than five pairs each of Griffon Vultures and Black Vultures and no more Lammergeiers breed in Bulgaria.     

Detrimental impacts of radiotransmitters on juvenile Louisiana Waterthrushes

ABSTRACT.   The Louisiana Waterthrush (waterthrush; Seiurus motacilla ) is a forest-dwelling, Nearctic-Neotropical migratory passerine that nests along streams. We attached radiotransmitters (0.6–0.8 g) to 12 nestling waterthrushes using snug, elastic loops. At three nests, adult waterthrushes were videotaped removing radio-tagged young from the nest. In addition, we recovered nine radio-backpacks (with two still attached to the carcasses of nestlings) near nests within a few days after attaching transmitters. Only one of 12 radio-tagged young was relocated more than 24 h after attaching the transmitter. Thus, the method of transmitter attachment we used was not effective. Using snug, nonelastic loops (e.g., nylon) for the harness may reduce the loss of transmitters, but may injure the skin as fledglings grow. Other possible alternatives include (1) gluing the transmitter to skin on the back of nestlings, (2) capturing fledglings in mist nets and attaching transmitters a week or more after fledging by which time contour feathers have grown and the likelihood of a parent removing the transmitter may be reduced, or (3) attempting to monitor fledglings without attaching transmitters. The success of the latter two alternatives would likely be enhanced by attaching transmitters to adults and then tracking them to locate their still-dependent fledglings.     

Brood parasitism increases provisioning rate, and reduces offspring recruitment and adult return rates, in a cowbird host

Interspecific brood parasitism in birds presents a special problem for the host because the parasitic offspring exploit their foster parents, causing them to invest more energy in their current reproductive effort. Nestling brown-headed cowbirds (Molothrus ater) are a burden to relatively small hosts and may reduce fledgling quality and adult survival. We documented food-provisioning rates of one small host, the prothonotary warbler (Protonotaria citrea), at broods that were similar in age (containing nestlings 8–9 days old), but that varied in composition (number of warbler and cowbird nestlings) and mass, and measured the effect of brood parasitism on offspring recruitment and adult returns in the host. The rate of food provisioning increased with brood mass, and males and females contributed equally to feeding nestlings. Controlling for brood mass, the provisioning rate was higher for nests with cowbirds than those without. Recruitment of warbler fledglings from unparasitized nests was 1.6 and 3.7 times higher than that of fledglings from nests containing one or two cowbirds, respectively. Returns of double-brooded adult male and female warblers decreased with an increase in the number of cowbirds raised, but the decrease was more pronounced in males. Reduced returns of warbler adults and recruitment of warbler fledglings with increased cowbird parasitism was likely a result of reduced survival. Cowbird parasitism increased the warblers’ investment in current reproductive effort, while exerting additional costs to current reproduction and residual reproductive value. Our study provides the strongest evidence to date for negative effects of cowbird parasitism on recruitment of host fledglings and survival of host adults.     

The adaptive function of construction of multiple non-breeding nests in birds

Several bird species construct multiple nests within a single breeding season that are not used for egg-laying. This behaviour has puzzled researchers for over 100 years, as nests are costly in time and energy to build, and there is no apparent adaptive function. We review the empirical evidence for several suggested non-exclusive functions and examine the plausibility of each. These functions are: (1) decoys to reduce predation or brood parasitism, (2) involvement in sexual selection and signalling of traits such as genetic quality or propensity for parental investment, (3) shelters for adults and/or fledglings, and (4) defence against nest usurpation. We argue that evidence for all these explanations is inconclusive, with correlational studies dominating the literature. We conclude that the idea that these nests evolved as decoys or as shelters for fledglings is unlikely and consider the most likely adaptive function to be signalling. Multiple nests are often involved in aspects of courtship, so are likely have some role in sexual selection in certain species. After pairing, individuals may be able to signal their parental abilities, inducing their partner to invest more in reproduction through differential allocation. We give suggestions for future work which we believe will advance our understanding of ancillary nest-building.     

Parent–offspring recognition in the Brown-headed Parrot Poicephalus cryptoxanthus

Recognition by vocal characteristics between parents and their offspring is thought to be ubiquitous in colonially nesting avian species. The Brown-headed Parrot Poicephalus cryptoxanthus nests in hollows in trees. However, when the chicks fledge they leave the nest and for the following three weeks spend their time in a tree where they are fed by their parents. As the fledglings are mobile and cryptic, returning parents must locate their own chicks. In this study a series of playback experiments was carried out, which showed that the chicks recognise their parents by voice. A simultaneous mirror experiment indicated that recognition was not reciprocated, although there may be alternative explanations for this behaviour. When the young began to forage with their parents, vocal stimuli did not induce any response from adult or chick. We suggest that vocal recognition becomes secondary to visual recognition as development proceeds.     

Odour-based natal nest recognition in the zebra finch (Taeniopygia guttata), a colony-breeding songbird

Passerine birds have an extensive repertoire of olfactory receptor genes. However, the circumstances in which passerine birds use olfactory signals are poorly understood. The aim of this study is to investigate whether olfactory cues play a role in natal nest recognition in fledged juvenile passerines. The natal nest provides fledglings with a safe place for sleeping and parental food provisioning. There is a particular demand in colony-breeding birds for fledglings to be able to identify their nests because many pairs breed close to each other. Olfactory orientation might thus be of special importance for the fledglings, because they do not have a visual representation of the nest site and its position in the colony when leaving the nest for the first time. We investigated the role of olfaction in nest recognition in zebra finches, which breed in dense colonies of up to 50 pairs. We performed odour preference tests, in which we offered zebra finch fledglings their own natal nest odour versus foreign nest odour. Zebra finch fledglings significantly preferred their own natal nest odour, indicating that fledglings of a colony breeding songbird may use olfactory cues for nest recognition.     

Social mating system and reproductive success in house wrens

Current models explaining the establishment and maintenanceof social monogamy and polygyny within avian populations typicallyassume that the reproductive success of polygynous males exceedsthat of monogamous males. This assumption is almost always supportedwhen the number of fledglings or recruits to future breedingpopulations is used to measure adult reproductive success. However,recent studies using DNA markers indicate that simple countsof fledglings or recruits may be a poor estimator of the numberof nestlings sired by the social father. In this paper, we comparethe number of genetic offspring produced by socially monogamousand polygynous house wren (TrogiodyUs atdon) males in nestsat which they were the social father. Polygynous males did,in fact, sire more nestlings in their own nests than did monogamousmales. Moreover, although we have not identified the sires ofextrapair nestlings, we document that even when all extrapairnestlings in this population are hypotheticaOy assigned to monogamousmales, die total reproductive success of polygynous males exceedsthat of monogamous males. These results and those of severalother recent studies are consistent with the assumption thatpolygynous males produce more offspring than monogamous males.     

A stab in the dark: chick killing by brood parasitic honeyguides

The most virulent avian brood parasites obligately kill host young soon after hatching, thus ensuring their monopoly of host parental care. While the host eviction behaviour of cuckoos (Cuculidae) is well documented, the host killing behaviour of honeyguide (Indicatoridae) chicks has been witnessed only once, 60 years ago, and never in situ in host nests. Here, we report from the Afrotropical greater honeyguide the first detailed observations of honeyguides killing host chicks with their specially adapted bill hooks, based on repeated video recordings (available in the electronic supplementary material). Adult greater honeyguides puncture host eggs when they lay their own, but in about half of host nests at least one host egg survived, precipitating chick killing by the honeyguide hatchling. Hosts always hatched after honeyguide chicks, and were killed within hours. Despite being blind and in total darkness, honeyguides attacked host young with sustained biting, grasping and shaking motions. Attack time of 1-5 min was sufficient to cause host death, which took from 9 min to over 7 h from first attack. Honeyguides also bit unhatched eggs and human hands, but only rarely bit the host parents feeding them.     

The role of parent–offspring interactions during and after fledging in the Black-legged Kittiwake

Most bird species endure a high mortality at fledging, and selection should favour parental behaviour diminishing these costs. Post-fledging parental care varies greatly among species and is often linked to parent–offspring recognition. In the Black-legged Kittiwake (Rissa tridactyla), fledglings need to return to the natal nest to be fed by their parents until independence. Rejections of fledglings by non-parent adults may be fairly violent, and parents are expected to recognize and help their chicks at the time of first return. However, previous cross-fostering experiments pointed out that parents are not able to recognize their chicks up to 15 days before fledging. In this paper, we study the behaviour of both parents and juveniles at fledging. We found that parents answered significantly more to their fledgling's calls than to those of others. Compared to silent juveniles, juveniles that called before landing were more likely to be accepted by their parents. No such pattern was observed with foreign juveniles, indicating that fledglings’ voice may carry individual identity. Furthermore, fledglings found their way back to the natal nest faster when parents attended the natal nest and reacted to their offspring's calls than when they were absent or inactive. Such interactions may therefore diminish juvenile mortality at fledging.