Intraflock variation in the speed of escape-flight response on attack by an avian predator

The benefits of flocking to prey species, whether through collective vigilance,dilution of risk, or predator confusion, depend on flock members respondingin a coordinated way to attack. We videotaped sparrowhawks attackingredshank flocks to determine if there were differences in thetiming of escape flights between flock members and the factorsthat might affect any differences. Sparrowhawks are surpriseshort-chase predators, so variation in the time taken to takeflight on attack is likely to be a good index of predation risk.Most birds in a flock flew within 0.25 s of the first bird flying,and all birds were flying within 0.7 s. Redshanks that werevigilant, that were closest to the approaching raptor, and thatwere close to their neighbors took flight earliest within aflock. Birds in larger flocks took longer, on average, to takeflight, measured from the time that the first bird in the flockflew. Most birds took flight immediately after near neighbors tookoff, but later flying birds were more likely to fly immediatelyafter more distant neighbors took flight. This result, alongwith the result that increased nearest neighbor distance increasedflight delay, suggests that most redshanks flew in responseto conspecifics flying. The results strongly suggest that thereis significant individual variation in predation risk withinflocks so that individuals within a flock will vary in benefitsthat they gain from flocking.     

Avoidance of headwinds or exploitation of ground effect—why do birds fly low?

ABSTRACT Birds often fly close to the ground or water. Wind shear theory predicts that wind speeds decline with proximity to the substratum, so birds might be expected to fly lower when flying upwind than when flying downwind. We tested this prediction and found that the wind shear equation is valid at heights below 4 m, with wind speed over a smooth surface ～40% lower at a height of 0.08 m than at 4 m. Birds that fly close enough to smooth substrata can also benefit energetically from ground effect, where vortices generated by their flight interact with the ground or water. This suggests that birds should use ground effect more when flying upwind than when flying downwind. We determined the percent time spent flying in ground effect by 21 species of passerine and non‐passerine birds flying in sheltered coastal aquatic and nearby terrestrial areas of County Cork, Ireland. We found that use of ground effect was uncommon for passerines, but common for a variety of non‐passerine waterbirds. However, phylogenetic analysis indicates no linkage between phylogeny and incidence of ground effect use and it is probable that incidence of use is determined by ecology rather than phylogeny. Great Cormorants (Phalacrocorax carbo) used ground effect most frequently over water (59.4% of time in flight). Over land, Barn Swallows (Hirundo rustica) used ground effect most often (19.8% of time). Phylogenetic contrasts regression analysis showed no significant relationship between use of ground effect and either wing aspect ratio or wing loading for 18 of our focal species, though simple linear regression analysis indicated that birds with greater wing loading used ground effect slightly (but significantly) more often. We found that 95% of Great Cormorants flying upwind used ground effect whereas only 35% did so when flying downwind. Few Black‐headed Gulls (Chroicocephalus ridibundus) used ground effect (probably because they fly high to locate prey), but still showed greater use when flying upwind (25%) than downwind (2.5%). When flying upwind in ground effect at the wind speeds encountered in our study, the velocity for minimum power (V_mp) for Great Cormorants was exceeded, suggesting theoretical benefits of about 14.3%. Our study indicates that several species exploit both wind shear and ground effect to minimize energy expenditure during commuting and foraging, but that others do not, because of either complexity of habitat morphology or the demands of their foraging ecology.     

SOARING BEHAVIOUR AND PERFORMANCE OF SOME EAST AFRICAN BIRDS, OBSERVED FROM A MOTOR-GLIDER

Various species of soaring birds were studied by following them in a motor-glider, mainly over the Serengeti National Park, Tanzania. The characteristics of thermal convection in the study area are described in general terms. The two vulture species of the genus Gyps live by scavenging among the herds of migratory ungulates, especially Wildebeest. They are not territorial, and gather in large numbers on kills. When raising young they may be obliged by game movements to forage at long distances from their nests. Their cross-country performance is adequate for a foraging radius of over 100 km in dry-season conditions. Their ability to compete with Spotted Hyaenas is thought to depend partly on this factor and partly on an advantage in arriving early at kills. These two species appear to find food more by watching other vultures than by searching for it directly. The Lappet-faced and White-headed Vultures are thought to be sedentary, and to depend on thorough searching of a fixed foraging territory, rather than on following migratory game. They have lower wing loadings than the Gyps vultures, and were not seen cross-country flying. They never gather in large numbers. The Hooded Vulture is a solitary nester, but it does fly across country, and does gather at kills. Vultures soar individually, and seem to be good at exploiting such phenomena as thermal streets. They do not travel in flocks. Tawny and Martial Eagles react positively to the glider, and are suspected of regarding it as potential prey. White Storks migrate between Europe and Africa, and also travel about within East Africa, by thermal soaring. They soar in flocks, and unlike vultures rely on co-ordinated social behaviour to locate thermals. In choosing their route, they often fail to react to obvious weather signs. They enter cumulus clouds from the bottom when thermalling, but probably do not climb far above cloudbase. Marabou Storks soar individually, but also sometimes travel in flocks. When doing so, they show less lateral spreading than White Storks, which reduces the effectiveness of the flock as a thermal-finding unit; on the other hand, they do seem to react to visible weather signs, like vultures or glider pilots. White Pelicans, which travel by thermal soaring between different lakes in the Rift Valley, show the most highly co-ordinated social soaring behaviour. Unlike White Storks, they fly in formation even when circling. Storks and pelicans showed more signs of alarm when approached by the glider than did the vultures or birds of prey. This could be due to their being preyed upon in flight, for instance by Martial Eagles. The basis of conventional thermal cross-country flying is outlined, and it is explained why the high wing loadings of the Gyps vultures are appropriate to their peripatetic habits. A method of thermal soaring without circling is discussed, and shown to be more readily feasible for small than for large birds. Some differences in soaring techniques between birds and glider pilots are interpreted in the light of this calculation. A case in which Black Kites apparently used this technique to soar in random turbulence is described. The cross-country speed attainable by thermal soaring should be similar to the cruising speed under power in both large and small birds. Rough calculations of the energy costs suggest that a large bird (White Stork) should reduce its fuel consumption by a factor of 23 by soaring rather than flying under power, whereas this factor would be only 2–4 for a small bird (Bonelli's Warbler). Other reasons why thermal soaring is an advantageous means of travel for large but not for small birds are also indicated.     

Do territorial and non‐breeding Australian Magpies Gymnorhina tibicen influence the local movements of rural birds in New Zealand?

Australian Magpies Gymnorhina tibicen sometimes attack and kill other birds in New Zealand. Here we assess how Australian Magpies influence the local distribution of other birds in New Zealand and identify the members of an Australian Magpie population that display the most agonistic acts. We conducted regular observations on six territorial breeding groups and three non‐breeding flocks of Australian Magpies for 1 year to determine (1) if other birds avoid flying or foraging close to Australian Magpies, (2) the proportion of passing birds that are attacked and (3) which social subunits of the Australian Magpie population are most aggressive. In comparison with adjacent Magpie‐free control areas, significantly fewer birds of a range of species (e.g. Common Blackbird Turdus merula, Skylark Alauda arvensis, Yellowhammer Emberiza citrinella) foraged close (i.e. ≤ 50 m) to both territorial breeding groups and non‐breeding Australian Magpie flocks; fewer birds were also recorded flying near (i.e. ≤ 50 m) territorial breeding groups but not non‐breeding flocks. Excluding Australasian Harriers (Circus approximans: see below), only 8% of birds recorded within 50 m of territorial breeding groups were observed being attacked. Attacks were most frequent when numerous birds occurred near Australian Magpies and species recorded in the highest frequencies were generally attacked most. Territorial breeding groups attacked 39% of passing Australasian Harriers. All attacks consisted of the victim being swooped at or chased; no physical contact was ever observed. Both adult male and female breeding Australian Magpies were seen attacking other birds; juveniles in breeding groups sometimes supported adults but never initiated attacks. Australian Magpies in non‐breeding flocks were not seen attacking other birds, except Australasian Harriers (attacked in 17% of appearances). Our results suggest that some birds avoid foraging and/or flying close to Australian Magpies because they are sometimes chased by breeding adults of both sexes; however, the proportion of passing birds actually attacked was small. The numerous published observations of Australian Magpie attacks are apparently biased heavily towards sensational events that are rare. Possible reasons why Australian Magpies attack other birds are discussed.     

Space use and agonistic behaviour in relation to sex composition in large flocks of laying hens

Some authors have found indications of subgroup formation when domestic fowl are forced to live together in large flocks, while others have not. In this study experiments were carried out to test the hypothesis that hens in large flocks have home ranges in parts of the pen and that they form subgroups. We also studied if this is influenced by males. In a tiered aviary system (density averaged 16 hens/m(2) of floor area) eight flocks of 568+/-59 ISA Brown laying hybrids were kept in pens. Half of the pens contained 1 male per on average 24 females (mixed flocks). At peak production (36-53 weeks of age) four females roosting closely together for about 14 days and four females roosting far apart from each other were taken out from each flock and put together in separate groups in small pens. Their agonistic behaviour was studied for 2 days before they were put back. This was repeated with new birds, resulting in 16 small sample groups being studied. At 70 weeks, three groups of 10 females per flock roosting closely together in different parts of the pen were dyed with different colours and their locations were observed for 2 nights and 2 days.The incidence of aggressive pecks during day 1 among birds that had been roosting close to each other tended to be lower (P=0.05) than among birds that had been roosting far apart. This effect was not significant among birds from all-female flocks, but among birds from mixed flocks (P<0.05). However, this indicates a recognition of roosting partners and possibly also a rebound effect of the males' reduction of female aggressiveness towards strangers. Irrespective of sex composition in the flocks, birds marked while roosting at the ends of the pens were significantly more often observed within these areas than in other areas of the pen during daytime and came back to the same roosting sites at night (P<0.05-P<0.001). This was not the case for birds from the middle of the pens, where the distribution in the pen in most cases did not differ from random. These results show that laying hens in large groups are rather constant in their use of space, which indicate the presence of home ranges. However, environmental features that facilitate localisation may be important. In summary, we think that these findings indicate the existence of subgroup formation.     

Experimental analysis of the social value of flocking by starlings (Sturnus vulgaris) in relation to predation and foraging

In groups of ten, indidual starlings, Sturnus vulgaris, spent significantly less time in surveillance than did individuals in smaller groups and responded more quickly than single birds to a flying model hawk. Captive starlings in flocks reduce their individual surveillance efforts, but their combined efforts still enable them to be more effective than single birds in the detection of predators. Foraging behaviour of flocks was observed by placing single starlings with groups of tricoloured blackbirds, Agelaius tricolor; the starlings reduced the time they devoted to surveillance at the same rate as if they were with other starlings.     

WHY FORAGING BIRDS IN TREES SHOULD CLIMB AND HOP UPWARDS RATHER THAN DOWNWARDS

This paper describes the energy cost of locomotion in birds foraging over vertical zones in trees. In particular, the energetically cheapest pattern for a bird flying among trees and moving within them is explored. For birds moving vertically by climbing and hopping (but not by flying) it should take less energy to climb and hop upwards in a tree and fly downwards to the next one, than to do the reverse. This is because part of the potential energy gained in climbing upwards may be used for subsequent horizontal progression to the next tree. For movements the other way, the potential energy is largely wasted during downward hopping and climbing within a tree. It is predicted that birds moving within trees by climbing and hopping (but not by flying) leave at a higher level than they arrive (whether the vertical movements within trees are along the trunk or among branches). These energetic considerations probably expose one selection pressure behind the morphology of the woodpecker—treecreeper type, which shows obvious adaptation for climbing upwards rather than downwards.     

Unique near isometric ontogeny in the pterosaur Rhamphorhynchus suggests hatchlings could fly

Rhamphorhynchus muensteri is one of the best‐known flying reptiles, represented by >130 well‐preserved fossil specimens, from hatchlings to full adults. The life history of this pterosaur remains controversial as to when in ontogeny they took flight. Here, we assess the growth of these animals based on the lengths of numerous key elements. We show that changes in the skeletal anatomy of this reptile across its post‐hatch size range reveal that R.muensteri exhibited overall near isometric growth in the wings, with slightly negative allometry in the humerus, radius and stronger negative allometry in the fourth metacarpal compared to body length, and slightly positive allometry in the second and third phalanges compared to body length. This pattern is near unique among flying vertebrates and suggests R.muensteri flew soon after hatching. In bats and birds, offspring do not typically fly until nearly adult sized. Conversely, near isometric growth in Rhamphorhynchus suggests it was a precocial flier and that individuals may have inhabited several sequential foraging niches over their lifespan, as some terrestrial and aquatic vertebrates do today.     

Metabolism during flight in two species of bats, Phyllostomus hastatus and Pteropus gouldii.

The energetic cost of flight in a wind-tunnel was measured at various combinations of speed and flight angle from two species of bats whose body masses differ by almost an order of magnitude. The highest mean metabolic rate per unit body mass measured from P. hastatus (mean body mass, 0.093 kg) was 130.4 Wkg-1, and that for P. gouldii (mean body mass, 0.78 kg) was 69.6 Wkg-1. These highest metabolic rates, recorded from flying bats, are essentially the same as those predicted for flying birds of the same body masses, but are from 2.5 to 3.0 times greater than the highest metabolic rates of which similar-size exercising terrestrial mammals appear capable. The lowest mean rate of energy utilization per unit body mass P. hastatus required to sustain level flight was 94.2 Wkg-1 and that for P. gouldii was 53.4 Wkg-1. These data from flying bats together with comparable data for flying birds all fall along a straight line when plotted on double logarithmic coordinates as a function of body mass. Such data show that even the lowest metabolic requirements of bats and birds during level flight are about twice the highest metabolic capabilities of similar-size terrestrial mammals. Flying bats share with flying birds the ability to move substantially greater distance per unit energy consumed than walking or running mammals. Calculations show that P. hastatus requires only one-sixth the energy to cover a given distance as does the same-size terrestrial mammal, while P. gouldii requires one-fourth the energy of the same-size terrestrial mammal. An empirically derived equation is presented which enables one to make estimates of the metabolic rates of bats and birds during level flight in nature from body mass data alone. Metabolic data obtained in this study are compared with predictions calculated from an avian flight theory.     

The flight speed of parent birds feeding young

I review previous models of the speed at which parent birds should fly when delivering food to their young. Norberg gives a graphical method of finding a parent's best flight speed. This speed maximizes the overall rate at which energy is delivered to the young. An alternative assumption is that a parent maximizes the net rate of delivery of energy. I suggest that in general we cannot distinguish between net rate and overall rate on the basis of whether the parent feeds itself. The best way to distinguish between these currencies may be to use qualitative predictions. I present new results on the effect of a constraint on energy expenditure on the parent's optimal speed. I show that the optimal speed when foraging should be less than the optimal speed when traveling. I also analyse the advantage to a parent of flying faster than the maximum range speed and evaluate previous empirical studies of the speed at which parent birds fly. Only one study claims that parent birds fly at the speed identified by Norberg, but I raise doubts about this claim.     

Optimal locomotion modes of foraging birds in trees

This paper compares the energy costs of various modes of locomotion of birds foraging in trees. For birds moving vertically in trees by climbing and hopping (but not by flying) the best choice of locomotion mode depends on the distance between visited trees in relation to the height h of the zone searched for food in trees.
When the distance between successively visisted trees is longer than about half the distance coverable in gliding flight with height loss h , then it is cheapest in energy to hop or climb upwards in a tree and fly downwards to the next tree. When the distance between successively visited trees is shorter than about half the distance coverable in gliding flight with height loss h , then it is cheapest in energy to move alternately downwards and upwards in trees (downwards in the first, upwards in the second, downwards in the third tree, etc.) and to fly level between trees.
Treecreepers and woodpeckers are adapted morphologically to the former mode, but more generalized tree foragers might use either mode depending on the spacing of trees.     

Recording raptor behavior on the wing via accelerometry

ABSTRACT.   Measuring body movements using accelerometry data loggers is a relatively new technique, the full applicability of which has yet to be tested on volant birds. Our study illustrates the potential of accelerometry for research on large birds by using the technique to record the behavior of three species of raptors, mainly during flight. A tri-axial accelerometer was deployed on a trained Harris' Hawk ( Parabuteo unicinctus ), Tawny Eagle ( Aquila rapax ), and Griffon Vulture ( Gyps fulvus ). Comparison of flight-related variables calculated from video footage and that estimated from the acceleration data showed that the latter provided considerable and accurate information (usually <10% error) about the behavior of the birds, including wing-beat frequency and when they glided and flapped. Acceleration data permitted tentative comparisons of relative movement-specific rates of energy expenditure for the Griffon Vulture flying up versus flying down a small hill. The accelerometry data appeared to suggest, as expected, that the Griffon Vulture expended more energy flying uphill than flying back down. Our preliminary findings indicate that studies using accelerometers can likely provide information about the detailed time–energy budgets of large birds. Such information would aid in comparative analyses of behavior and energetics, and may also enhance efforts to conserve declining bird populations.     

Familiarity Breeds Tolerance: the Development of Social Stability in Flocking Siskins (Carduelis spinus)

Behavioural integration associated with the fusion of two flocks is analyzed in captive siskins (Carduelis spinus) by quantifying changes in social behaviour with time since joining. In general there was an increase in the incidence of tolerant behaviour, supplanting attacks and hopping withdrawals with time since fusion of the flocks. However, the number of displays and flights showed the opposite, negative, trend. Taking dominance status into account, the greatest change in behaviour with time since joining is an increase in tolerance by dominants of new flock companions. Factorial analysis of correspondences was used to study how different birds changed their behaviour with time since joining a flock. This analysis showed that the introduction of new birds did not disrupt relationships with familiar birds, and that residents are dominant in interactions with the incoming new flock companions. The analysis also demonstrated that relationships within the new flock had stabilized 20 days after the flocks had joined. The characteristics of the socially integrated group of siskins are quite similar to those described by Rohwer & Ewald (1981) in their shepherds hypothesis: dominants tolerate their subordinates feeding in close proximity, offering them a profitable feeding area, but also supplant them to obtain food; both dominants and subordinates benefit from being in a flock. As a consequence, constant changes of membership in flocks is costly not only because birds lose dominance status, but also the advantages of clear dominant and subordinate roles.     

Hovering and intermittent flight in birds

Two styles of bird locomotion, hovering and intermittent flight, have great potential to inform future development of autonomous flying vehicles. Hummingbirds are the smallest flying vertebrates, and they are the only birds that can sustain hovering. Their ability to hover is due to their small size, high wingbeat frequency, relatively large margin of mass-specific power available for flight and a suite of anatomical features that include proportionally massive major flight muscles (pectoralis and supracoracoideus) and wing anatomy that enables them to leave their wings extended yet turned over (supinated) during upstroke so that they can generate lift to support their weight. Hummingbirds generate three times more lift during downstroke compared with upstroke, with the disparity due to wing twist during upstroke. Much like insects, hummingbirds exploit unsteady mechanisms during hovering including delayed stall during wing translation that is manifest as a leading-edge vortex (LEV) on the wing and rotational circulation at the end of each half stroke. Intermittent flight is common in small- and medium-sized birds and consists of pauses during which the wings are flexed (bound) or extended (glide). Flap-bounding appears to be an energy-saving style when flying relatively fast, with the production of lift by the body and tail critical to this saving. Flap-gliding is thought to be less costly than continuous flapping during flight at most speeds. Some species are known to shift from flap-gliding at slow speeds to flap-bounding at fast speeds, but there is an upper size limit for the ability to bound (~0.3 kg) and small birds with rounded wings do not use intermittent glides.     

Downsized Dinosaurs: The Evolutionary Transition to Modern Birds

Living birds are the most diverse land vertebrates and the heirs of a rich chapter in the evolution of life. The origin of modern birds from animals similar to Tyrannosaurus rex is among the most remarkable examples of an evolutionary transition. A wealth of recently discovered fossils has finally settled the century-old controversy about the origin of birds and it has made the evolutionary saga toward modern birds one of the best documented transitions in the history of life. This paper reviews the evidence in support of the origin of birds from meat-eating dinosaurs, and it highlights the array of fossils that connect these fearsome animals with those that fly all around us.     

Social dominance in free-living Willow Tits Parus montanus: determinants and some implications of hierarchy

Several features of social dominance among Willow Tit Parus montanus winter flocks were examined during a four-winter study. Birds of both sexes were evenly distributed over the 33 flocks studied. In nearly half of the flocks there was an adult pair accompanied by yearlings, but one-third of the flocks consisted of more than two adults with yearlings. Males were found to be dominant over and larger than females. Within a sex, yearlings were usually subordinate to adults. The effect of size on dominance, after controlled for sex and age, remained obscure in our field data. The hierarchical status of an individual was found to rise or at least stay the same in different years, which supports the "hopeful dominants" hypothesis, i.e. birds stay in flocks hoping to achieve a higher status in the future. The ranks of mates correlated highly significantly, implying that high-ranking birds were paired with other high-ranking birds and low-ranking birds with other low-ranking birds. Birds of different age and sex did not show any differences in the proportion of initiated aggressive encounters directed at other individuals. However, males were more aggressive to other males than to females and also tended to behave less aggressively towards their own mates than towards other individuals in the flock. This could be a male strategy to strengthen the pair-bond and to enhance mate protection described earlier in     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

ASPECTS OF THE BEHAVIOUR AND ECOLOGY OF MIXED-SPECIES BIRD FLOCKS IN KASHMIR

Mixed-species flocks of birds were observed between the end of July and late August, principally at Daksum, Kashmir, 2250 m. The species composition and the numbers of individuals in flocks changed during this period; these changes are attributed to resident territory holders and migrant birds joining the flocks. Within the flock different species showed some differences in foraging stations, but nevertheless often appeared to be taking the same type of food. Participant species had different roles in the flock organization. Behaviours involving the entire mixed-flock acting as a unit included path reversal after encounters with avian predators and a tendency to follow set routes. The mixed- species flock exerted an attractive influence on aggregations of species not normally participant.
Similarities between the flocks described in this study and those recorded by other workers are discussed. While different species may derive different benefits from joining these flocks, advantages that could benefit some or all participants include the receipt of information on good feeding areas in an unfamiliar locality, the avoidance of time wasted on feeding on substrates which have been very recently harvested, the beating effect to increase prey availability, and enhanced safety from predators, perhaps through differential alertness of different species and specialized anti-predator behaviour.     

The diet of the New Holland honeyeater,Phylidonyris novaehollandiae

New Holland honeyeaters collect nectar, manna or honeydew for energy and hawk small flying insects for protein. The insects taken were usually Diptera and Hymenoptera weighing 0.7 mg dry weight or less. Net rates of energy gain from hawking small flying insects were usually less than 20 J min^?1 and sometimes negative and insufficient to meet the bird's daily energy requirements. Those from feeding on nectar, manna or honeydew were usually above 40J min^?1 and often above 400J min^?1 at dawn and the birds depended on these carbohydrates for energy. Nectar, manna and honeydew contained negligible amounts of protein, and the birds used small flying insects as sources of protein, and presumably other nutrients. Given that carbohydrate resources supply better rates of energy gain than insects. New Holland honeyeaters should collect their energy requirements from carbohydrates and only collect sufficient insects to satisfy their protein requirements. Estimates of the food intakes of both non-breeding and breedig birds showed that they did this. Non-breeding New Holland honeyeaters collected from 72 to 125 (mean 92) kJ of carbohydrates per day and 17 to 58 (mean 31) mg of protein per day. These meet the daily energy (75 kJ) and protein (20 mg) requirements of the birds. Breedig birds collected more carbohydrates and more insects, but in proportion to their increased energy and protein requirements respectively. New Holland honeyeaters are probably limited by their ability to meet their energy requirements from nectar, manna or honeydew and not by insects. Non-breeding birds collected their protein requirements in about 10 min of insect-feeding, but spent from 33 to 90% of the day collecting carbohydrates to meet their energy requirements. The maintenance requirement of 20 mg of protein per day for New Holland honeyeaters is about 25% of that estimated from standard equations for a bird of the same size. This low level may have evolved in response to low energy availability.     

Do bird captures reflect migration intensity? – Trapping numbers on an Alpine pass compared with radar counts

A limitation of standardized mist netting for monitoring migration is caused by the lack of knowledge about the relationship between trapped birds and birds flying aloft. Earlier studies related nocturnal radar counts with trapping data of the following day. In this study, we compared for the first time data gathered simultaneously by radar and mist netting, separately for diurnal and nocturnal migration. Trapping numbers were strongly correlated with migratory intensities measured by radar (r>0.6). A multiple regression analysis, including wind speed and wind direction explained 61% of variation in the number of captures. During the night, and particularly with favourable winds, birds flew at higher altitudes and hence escaped the nets to a higher proportion. The number of nocturnal migrants trapped during daytime was well correlated with migratory intensities observed by radar in the preceding night. The diurnal time patterns, however, revealed fundamental differences between trapping counts and radar observations. This was mainly due to increasing and decreasing flight altitudes in the course of the night, and by the limitations of the radar technique that underestimates migratory intensities during the day when birds aggregate in flocks. In relation to the migratory intensity recorded by radar, diurnal migrants are trapped in a much higher proportion than nocturnal migrants. Finally, our results confirm that trapping data from a site hardly used for stopover are well suited to represent the ongoing migration during the day and night.