Central place foraging in the white-fronted bee-eater

Orians & Pearson (1979) proposed a series of models of optimal central place foraging. I tested their model for single-prey loading species (those that carry items singly) with a natural population of white-fronted bee-eaters (Merops bullockoides) in Kenya. These insectivorous birds breed in colonies but defend nearby foraging territories from which they gather food for their nestlings. Qualitatively, two predictions of the model were upheld: as travel time increased, birds carried larger insects to their nests and spent longer periods of time foraging. Also, as distance increased, birds were increasingly selective of which insects they ate versus carried to the colony. Quantitative comparisons were also made using observed travel times and rates of insect capture. As distance increased, (1) the observed sizes of insects carried approached the values predicted by the Orians-Pearson model, (2) observed foraging times diverged from the predicted values, and (3) birds spent more time foraging between round trips in a relatively unprofitable area near the colony. These results are examined in light of the theory of central place foraging.     

The effect of prey abundance and foraging tactics on the population dynamics of a social, territorial carnivore, the spotted hyena

We used naturally occurring spatial and temporal changes in prey abundance to investigate whether the foraging behavior of a social, territorial carnivore, the spotted hyena ( Crocuta crocuta ), conformed to predictions derived from the ideal free distribution. We demonstrate that hyenas in the Ngorongoro Crater, Tanzania, redistributed themselves from less profitable to more profitable areas, even when this required them to undertake foraging trips to areas beyond their clan territory boundary, or required normally philopatric females to emigrate. As expected for a system with rank related access to food resources in the territory, females of low social status foraged more often outside their territory and were more likely to emigrate than dominant females. Probably because Crater hyenas regularly foraged outside their territories, there was no correlation between clan size and prey density within territories, suggesting that clan sizes may have exceeded the carrying capacity of territories. A substantial decline of the hyena population in the Crater from 385 adults in the mid 1960s to 117 in 1996 was most likely due to a substantial decline of their main prey. The decline in the hyena population was associated with a decline in the size of clans but not in the number of clans. The number of clans probably remained constant due to emigration by females from large clans into vacant areas or clans with no adult females, and because hyenas regularly fed in areas containing concentrations of prey beyond their territory boundary. Between 1996 and 2003 annual recruitment rates of Crater hyenas consistently exceeded annual mortality rates, resulting in an almost doubling of the adult population. This increase was most likely due to an increase in prey abundance, a relatively low level of predation on hyenas by lions ( Panthera leo ), and an absence of high levels of disease related mortality.     

THE RECOGNITION AND TREATMENT OF VENOMOUS AND NON-VENOMOUS INSECTS BY SMALL BEE-EATERS

The manner in which bee-eaters cope with the stinging honey-bees which comprise their principal food was investigated by aviary studies of Merops bullocki. Worker Apis are de-venomed by an innate sequence of beating the insect and rubbing its abdomen against the perch. Treatment improves with experience. Drone bees are distinguished by the birds from workers but are not instantly recognised as non-venomous; they are subjected to low intensity rubbing combined with non-venomous prey treatment concomitant with their size. Non-Hymenoptera, including some insects which superficially resemble bees, are recognised as non-venomous and are not rubbed, but are beaten until immobilized. Bee-eaters appear to have partial immunity to venom.     

Space and resource use by brown hyenas Hyaena brunnea in the Namib Desert

Two clans of brown hyenas were studied on the arid chilly Namib Desert coast where very large seal colonies are present. Carcasses from seals were abundant during summer and autumn following the seal pupping season, and scarce in winter and spring. Hyenas fed predominantly on carcasses. Members of one clan of three hyenas obtained most of their food along 3 km of coastline adjacent to the seal colonies. The other clan of nine individuals obtained food from an area along 6 km of coastline north of the seal colonies. Although carcasses were usually scavenged, seal pups were seen to be killed on occasions. Time spent foraging was not affected by carcass availability. Territories of the two clans overlapped and individual home-range size varied from 31.9 to 220 km². Food availability apparently had no influence on foraging activities but the dispersal of food affected territory and group sizes. Over abundance of food results in the maintenance of territories extending beyond the distribution of food. The shape and area of these territories may result from the cultural inheritance of space.     

The social organization of non-breeding Magpies Pica pica

The social behaviour of non-breeding individuals in a colour-marked population of Magpies was studied. In early autumn most non-breeders began to forage in a common area, the 'Non-breeding Flock Area'. A few individuals remained on their natal territories away from the flock area as solitary non-breeders. Most non-breeders were first-year birds, but some were second years or adults which no longer held a territory. Birds foraged in groups, their food intake rate differing with both group size and location. It appeared that the area in which birds foraged had a significant effect on food intake rate; foraging groups tended to form at sites rich in food. There was a dominance hierarchy amongst non-breeders; an individual's foraging behaviour, survivorship and chances of breeding were status dependent, with subordinates feeding less in groups and being less likely to survive and breed. Solitary individuals' chances of breeding were similar to those of high status birds, although their survivorship to breeding age may have been lower. Non-breeding Magpies are compared with non-breeders of other species and the factors which may influence their social behaviour are discussed. It is suggested that remaining as a solitary non-breeder is a viable alternative to becoming a low-status flock member for some birds.     

Aggressiveness in king penguins in relation to reproductive status and territory location

King penguins, Aptenodytes patagonicus, vigorously defend small territories in very dense colonies. The egg-laying season lasts approximately 4 months, but only pairs that reproduce during the first half of the period succeed in fledging a chick. I examined various factors affecting aggressiveness of king penguins during the breeding season and focused on the differences between central and peripheral territories. Pairs on peripheral territories experienced twice as many encounters with avian predators as did central birds. The vast majority of peripheral birds were late breeders, indicating that reproductive success was very low among penguins defending a territory on the edge of the colony. Time invested in territory defence and rate of agonistic encounters between breeding neighbours increased from the incubation to the brooding period. Parents gave most threat displays to territorial neighbours when the chick was very young and just before crèche formation. Distance to colony edge was not related to aggressiveness in incubating birds, however, the rate of pecking and flipper blows increased from the edge to the centre during brooding. In addition, aggressiveness of breeding penguins towards travelling birds trespassing into their territory increased with distance to edge. Early breeders were not more aggressive than late breeders but the proportion of time spent in territory defence increased with the number of days a bird had spent incubating. As expected, I did not detect any sex difference in aggressive behaviour. Birds occupying territories on the beach were generally more aggressive during the incubation period than those located on the valley sides. Reproductive status (incubating versus brooding) and territory location were the main factors explaining the various levels of aggressiveness observed in breeding king penguins. Copyright 2000 The Association for the Study of Animal Behaviour.     

The discriminant power of biometrics for sex determination in European Bee-eaters Merops apiaster

ABSTRACT

Capsule A combination of several biometric measures enables the reliable sexing of the European Bee-eater Merops apiaster, a species with subtle sex differences in plumage and morphometry.

Aims To explore variation in biometrics and their suitability to discriminate sex in adult European Bee-eaters Merops apiaster.

Methods We sampled populations of a long-distance migratory species with low sexual size dimorphism, the European Bee-eater Merops apiaster, from colonies in western Iberia (Portugal) and Central Europe (Germany) to investigate body size variation and derive population-specific and general sex discriminant functions.

Results Overall, male Bee-eaters were larger than females while Bee-eaters from Germany and Portugal did not differ in size, except for primary length and tail length. The best single measurement to discriminate sex was wing length for Portuguese birds and tail tip length for German birds, as in the combined dataset. Multivariate discriminant functions of head-bill, wing and tail tip lengths provided the highest discriminant accuracy, discriminating sex for 91% of the birds from Portugal, 96% from Germany and 94% when using the combined dataset. Nonetheless, the discriminant accuracy remained high in the functions using only two variables for Portugal (head-bill and tail tip 91%, head-bill and wing 88%, wing and tail tip 88%), Germany (head-bill and tail tip 95%, wing and tail tip 97%) and in the combined dataset (head-bill and tail tip 92%, wing and tail tip 93%).

Conclusions Population specific discriminant functions allow sexing of European Bee-eaters by morphometry with high degree of accuracy at least for Iberia and Central European populations. Such discriminant functions can be used to assign the sex of adult Bee-eaters reliably and swiftly while the bird is still in the hand, highlighting the potential of these functions for rapidly sexing species with low degrees of sexual size and plumage dimorphism.     

Partial migration in inexperienced pied avocets Recurvirostra avosetta: distribution pattern and correlates

Birds exhibit a range of wintering behaviour from strictly migrant to strictly resident species. In partially migrating ones, some birds overwinter within their breeding region (resident birds) while others, although breeding in the same area, winter far away (migrant birds). Accordingly, choosing a wintering region is a key stage in the annual life cycle of birds, notably for inexperienced first‐year individuals. The present study aimed to investigate this issue, and more specifically to study the distribution pattern during winter and factors influencing the wintering behaviour of first‐year pied avocets Recurvirostra avosetta. Based on a 10‐yr ringing study carried out on five of the major French breeding colonies distributed along the Atlantic coast, we showed the coexistence of different wintering tactics. The resident tactic was predominant (approximately 86% of the 575 birds re‐sighted), while the other birds adopted migration. Among resident individuals, two different tactics occurred: 43% of birds overwintered within their natal colony, whereas the others wintered in another site located at relatively close proximity along the French Atlantic coast. Hatching date was a consistent predictor of all wintering tactics. More specifically, the probability of migrating was the highest for early‐hatched birds, and for resident ones, the probability of wintering within their natal colony rather than in another French site was the highest for both median‐ and late‐hatched individuals. In addition, a colony effect was demonstrated for resident birds. Several biological interpretations, including social system, variations in both individual body condition and habitat quality, were put forward to explain these correlates.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

Stepping stones of life: natal dispersal in the group-living but noncooperative speckled warbler

In most cooperatively breeding birds the offspring of one sex, usually male, delays dispersal to remain on the natal territory and helps its parents to rear subsequent young. Thus delayed dispersal could be the first step in the evolution of cooperative breeding. We studied natal dispersal in a population of the group-living speckled warbler, Chthonicola sagittata, based on observations of a colour-banded population over 3 years. Unlike other group-living members of the Acanthizinae, all juvenile males in this population dispersed to settle on foreign territories as subordinates, which do not help rear the young. Speckled warblers showed all the life history traits that are thought to result in a saturated habitat and lead to delayed dispersal: they were sedentary, had high adult survival and had a male-biased sex ratio. However, they differed from other acanthizids in occurring at low density (0.18 birds/ha) on large breeding territories (6-12 ha), with a maximum of two males per territory. This may allow subordinates to live on foreign territories yet avoid aggression from dominants. A benefit of dispersal is that it provides an additional route to gaining a breeding vacancy. Dispersers can acquire vacancies on their new territory or on a neighbour's, but incest avoidance would be likely to constrain nondispersing males to neighbours' territories. A model of relative lifetime success showed that the survival benefits of natal philopatry are unlikely to outweigh this benefit of dispersal.     

Floater interference reflects territory quality in the Spanish Imperial Eagle Aquila adalberti: a test of a density‐dependent mechanism

We report on an 11‐year study of floater interference in a population of Spanish Imperial Eagles Aquila adalberti. We analysed changes over the years in the productivity of 15 territories to test predictions of two hypotheses of density‐dependent productivity in relation to the presence of floaters (birds without territories). According to the ‘interference' hypothesis, the frequency of intrusion by floaters increases with density, resulting in a decrease in productivity. Thus, in a high‐density population a negative relationship between floater intrusions and productivity of the territory is expected. In contrast, under the ‘habitat heterogeneity' hypothesis, as density increases a higher proportion of individuals is forced to occupy lower quality habitats. Support of this hypothesis requires that floaters detect differences in quality among territories and preferentially visit the better quality territories. Consequently, a positive relationship between floater intrusions and productivity is expected. Results showed that floaters tended to visit their natal area at the beginning of the breeding season. Among floater Eagles, males made significantly more intrusions per day than did females, but females stayed in the natal population for longer each year than males. Floater intrusions and productivity were highly positively correlated, supporting the ‘habitat heterogeneity' hypothesis; individuals were apparently able to assess the quality of a territory and, at the frequencies observed, their interference with the breeding pair had no obvious negative effect on productivity.     

Assimilation efficiency and growth of nestling insectivores

The efficiency of energy assimilation by Pacific Swallows Hirundo tahitica and Blue-throated Bee-eaters Merops viridis was measured in captive nestlings fed on their natural diets. Assimilation efficiency (AE) was 68% in Pacific Swallows, similar to published values for altricial nestlings, including other hirundines. AE was unusually low in Blue-throated Bee-eaters, conforming to results with the European Bee-eater Merops apiaster , at around 57%. As expected, nestling growth was positively correlated with daily metabolized energy. Estimates of daily metabolized energy for captive reared nestlings, closely resembled estimates obtained for wild youngsters in an earlier study using different methods.     

Fine‐scale habitat use during the non‐breeding season suggests that winter habitat does not limit breeding populations of a declining long‐distance Palearctic migrant

For migrant birds, which habitats are suitable during the non‐breeding season influences habitat availability, population resilience to habitat loss, and ultimately survival. Consequently, habitat preferences during winter and whether habitat segregation according to age and sex occurs directly influences migration ecology, survival and breeding success. We tested the fine‐scale habitat preferences of a declining Palearctic migrant, the whinchat Saxicola rubetra, on its wintering grounds in west Africa. We explored the influence of habitat at the territory‐scale and whether dominance‐based habitat occupancy occurs by describing the variation in habitat characteristics across wintering territories, the degree of habitat change within territories held throughout winter, and whether habitat characteristics influenced territory size and space‐use within territories or differed with age and sex. Habitat characteristics varied substantially across territories and birds maintained the same territories even though habitat changed significantly throughout winter. We found no evidence of dominance‐based habitat occupancy; instead, territories were smaller if they contained more perching shrubs or maize crops, and areas with more perching shrubs were used more often within territories, likely because perches are important for foraging and territory defence. Our findings suggest that whinchats have non‐specialised habitat requirements within their wintering habitat of open savannah and farmland, and respond to habitat variation by adjusting territory size and space‐use within their territories instead of competing with conspecifics. Whinchats show a tolerance for human‐modified habitats and results support previous findings that some crop types may provide high‐quality wintering habitat by increasing perch density and foraging opportunities. By having non‐specialised requirements within broad winter habitat types, migrants are likely to be flexible to changing wintering conditions in Africa, both within and across winters, so possibly engendering some resilience to the rapid anthropogenic habitat degradation occurring throughout their wintering range.     

Experimentally increased food resources in the natal territory promote offspring philopatry and helping in cooperatively breeding carrion crows

Kin-based societies, where families represent the basic social unit, occur in a relatively small number of vertebrate species. In the majority of avian kin societies, families form when offspring prolong their association with the parents on the natal territory. Therefore, the key to understanding the evolution of families in birds is to understand natal philopatry (i.e. the tendency to remain on the natal territory). It has been shown that, within populations, the strength of the association between parents and offspring (i.e. family stability) increases when offspring dispersal is constrained by external environmental factors, but it is unclear whether and how family wealth influences juvenile dispersal decisions. Here, we show that young carrion crows (Corvus corone corone) from territories that were food-supplemented year-round were more philopatric and more likely to help at their family's nest than the unfed ones. The results suggest that offspring philopatry and helping behaviour are influenced by the quality of 'home' and that the availability of food resources positively affects the cohesion of the family.     

Daughters on request: about helpers and egg sexes in the Seychelles warbler

The Seychelles warbler (Acrocephalus sechellensis) was an endangered endemic of the Seychelles islands where, until 1988, the entire population of ca. 320 birds was restricted to the one island of Cousin Island (29 ha). Additional breeding populations were successfully established on the islands of Aride (68 ha, 1988) and Cousine (26 ha, 1990) and now with the existence of ca. 2000 warblers on three islands the conservation status of the warbler has improved from endangered to vulnerable. Emigration from the island is extremely rare, so birds that disappeared were known to have died. Almost every bird on Cousin Island has been individually colour ringed and monitored throughout all breeding attempts during a 17-year period (1985-2002; total ca. 2400 birds). These birds were also blood sampled for molecular parentage and sex analyses. Therefore the lifetime reproductive success of many birds is known. Although warblers can breed independently in their first year, some individuals remain in their natal territory as subordinates, and often help by providing nourishment to non-descendant offspring. The frequency of 'helping' is affected by habitat saturation and variation in territory quality (insect prey availability). The long-term benefits of helping are higher for daughters than for sons, and it is therefore no wonder that most helpers are daughters from previous broods. Furthermore, on low-quality territories breeding pairs raising sons gain higher fitness benefits than by raising daughters, and vice versa on high-quality territories. Female breeders adaptively modify the sex of their single-egg clutches according to territory quality: male eggs on low quality and female eggs on high quality. The Seychelles warbler is a beautiful example of behavioural and life-history adaptations to restricted circumstances.     

Soil penetrability as a key factor affecting the nesting of burrowing birds

Soil penetrability resistance was found to be crucial for nest site selection of all three Central European burrowing bird species—Sand Martins (Riparia riparia), European Bee-eaters (Merops apiaster), and Eurasian Kingfishers (Alcedo atthis). Soil penetrability resistance measurements were used to find out whether increased hardness of unexcavated banks is the key factor affecting the presence of burrowing birds. All three species avoided banks composed of too compact or too loose soils. Birds discriminated not only between high- and low-quality breeding banks, but also between different soil strata within banks. In banks with generally low penetrability resistance, Sand Martins preferred soil strata with the highest available penetrability resistance and compactness to avoid hole collapses. There was a preference for hard soil below holes to serve as a resistant platform when birds begin to dig their holes. In Sand Martins, the penetrability resistance level affected physical characteristics of holes such as tunnel length and dimensions of the orifices. Excessive compactness—and probably not high talus presence—was a major cause of abandonment of Sand Martin localities. A high penetrability resistance is the crucial factor for site selection in Sand Martins.     

栗喉蜂虎捕食差异

2010年4～5月和2011年4～6月,用焦点动物取样法对云南省保山市潞江镇怒江河谷栗喉蜂虎捕食差异进行了观察。对栗喉蜂虎雌性成年鸟、雄性成年鸟、青年鸟捕食种类、数量、成功率、食物组成、食物资源量、处理食物成功率及处理时间进行了统计。结果表明,栗喉蜂虎捕食种类包括7类昆虫,以膜翅目Hymenoptera、蜻蜓目Odonate、鳞翅目Lepidoptera昆虫为主;成年雄鸟捕食频次高于成年雌鸟和青年鸟;成年鸟与青年鸟处理食物成功率无显著差异,但二者捕食成功率和处理食物时间差异显著。     

Seasonal and Diurnal Changes in Aggression and Territory Size in the Black-headed Gull (Larus ridibundus L.) on Islands in the Middle Reaches of the Vistula River

Seasonal and diurnal changes in territory size and frequency of aggressive interactions of various intensity were examined in colonies of the black-headed gull (Larus ridibundus) on river islands and ponds of central Poland. The highest frequency of the total aggression and the largest territories were recorded before the egg-laying stage, this being related to the establishing of territories, defence of the mate, and defence of nest material. Later in the season both aggression and territory size declined. Unlike the large species of gulls, black-headed gulls did not increase their aggression and territory size during hatching of the young. High-intensity aggressive behaviours (choking, attacking and fighting) were not so frequently displayed as low-intensity aggressive postures (upright, long call, forward) in each phase of the breeding cycle. Presumably this was because the former were more costly in terms of energy. Peak territorial activity occurred in the late afternoon. An increase in the frequency of intense aggressive behaviours (attack, fight) at that time was combined with an increased mobility of birds (departures to and arrivals from foraging sites). A reduced frequency of high-intensity aggressive displays in the period when the largest number of birds was present in the colony was likely to be adaptive.     

How does social behavior differ among sperm whale clans?

When individuals primarily associate with and learn from those who behave similarly, society and culture become closely tied. Sperm whales (Physeter macrocephalus) exhibit multilevel social structure, the levels of which are differentiated in part by characteristic cultural behaviors. Sperm whales are organized into sympatric clans, with distinctive vocal repertoires that are socially learned. Other behaviors, such as movement patterns and foraging, also differ among clans. Here we ask whether the clan partition also includes divergences in social behavior. Off the Galápagos Islands, members of two clans differed consistently in diving synchrony, heterogeneity, and temporal stability of social relationships. While number of associates (indicated by social unit, group, and cluster sizes) were similar between clans, Regular clan members dived more synchronously and had more homogeneous relationships than the Plus‐One clan members. Plus‐One social units had generally longer associations than those of the Regular clan. Differences in surface‐time coordination and quality of social relationships are likely byproducts of the clan segregation, which could affect alloparental care giving, therefore scaling up to differential calf survival rates between clans. This new dimension of behavioral divergence between sperm whale clans indicates that sympatric, socio‐cultural entities of nonhumans can also display characteristic social behavior.     

Life history of hamadryas baboons: Physical development,infant mortality,reproductive parameters and family relationships

Demographic and life history parameters were estimated for a band of free-ranging hamadryas baboons, observed for 5.5 years in Ethiopia. Age-related changes in body weight and dentition were found to be delayed relative to laboratory-reared baboons. On the average, females reached menarche at 4.3 years of age and had their first infant at the age of 6.1 years. The mean interbirth interval was 24 months if the infant survived this period. The survival of infants and juveniles was higher compared to Amboseli yellow baboons, but somewhat lower than in gelada baboons in the Simen Mountains. Males acquired their first juvenile or adult female at the age of 8.5 to 11 years. Male-female pair-bonds lasted several years in most cases. The Cone Rock baboons were organized in a four-level social structure. The troop could split into bands, bands were divided into clans, and clans into one-male units with bachelor followers. The exchange of individuals between social units predominantly occurred within the band. All males of known origin became adult members of their presumed natal clan. Most females transferred also within clans, and juvenile females tended to remain in their natal clan. Females lost by one male to several rivals tended to reassemble in the same new one-male units later on.