Sunbird hovering behavior is determined by both the forager and resource plant

The long‐standing paradigm that pollination systems adapted to hovering birds evolved only in the New World was recently challenged by the discovery of hovering pollination by Old World specialized passerine pollinators. This raises the possibility that hovering pollination may evolve more easily than previously believed, given sufficient selective pressure on plant traits, on nectarivory, or both. We observed foraging behavior by the sunbird Cyanomitra oritis at flowers of the native Old World plant Impatiens sakeriana. We measured the length of pedicels and peduncles (PedPed length), which can make the flowers difficult to reach while the bird perches on the stem, and determined if it influenced sunbird hovering or perching at a flower. Detailed analyses of video recordings showed that sunbirds only hovered at flowers with a long PedPed, whereas they employed both foraging modes when an adequate perch was available. A hovering sunbird could deplete nectar in a shorter time than a perching one. The frequency of visits was not greater at flowers with longer PedPed or with more open I. sakeriana flowers in the vicinity. Our study provides evidence that sunbird behavior does not follow simple energetic models, and that some sunbird pollination systems in the Old World resemble highly specialized hummingbird systems in the New World much more than expected, especially the overall adaptation of the system to bird hovering.     

Pollination of the understorey palm Calyptrogyne ghiesbreghtiana by hovering and perching bats

Inflorescences of the Central American understorey palm Calyptrogyne ghiesbreghtiana offer bats fruit-like flower tissue as reward for visitation. Bats visited the inflorescences in two different modes: frugivores (Phyllostomidae: Stenodermatinae, Carolliinae) foraged on the inflorescence while perched, whereas bats that were primarily nectarivores (Phyllostomidae: Glossophaginae) also consumed flower tissue, but did so while hovering. Fruit set was significantly lower in inflorescences that had received only hovering visits, indicating that perching behaviour offers better possibilities for pollen transfer. The pollination system of C. ghiesbreghtiana is probably adapted to perching frugivorous bats with nectar-specialist Glossophagines as non-optimal participants. Seen in an evolutionary perspective the C. ghiesbreghtiana– glossophagine bat interactions may demonstrate recruitment of new pollinators from non-destructive visitors.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 281–288.     

Design and Experiment of a Deformable Bird-inspired UAV Perching Mechanism

Energy consumption and acoustic noise can be significantly reduced through perching in the sustained flights of small Unmanned Aerial Vehicles(UAVs).However,the existing flying perching robots lack good adaptability or loading capac-ity in unstructured environments.Aiming at solving these problems,a deformable UAV perching mechanism with strong adaptability and high loading capacity,which is inspired by the structure and movements of birds'feet,is presented in this paper.Three elastic toes,an inverted crank slider mechanism used to realize the opening and closing movements,and a gear mechanism used to deform between two configurations are included in this mechanism.With experiments on its performance towards different objects,Results show that it can perch on various objects reliably,and its payload is more than 15 times its weight.By integrating it with a quadcopter,it can perch on different types of targets in outdoor environments,such as tree branches,cables,eaves,and spherical lamps.In addition,the energy consumption of the UAV perching system when perching on objects can be reduced to 0.015 times that of hovering.     

Why do some nectar foragers perch and others hover while probing flowers?

Diurnal hawkmoths, Hemaris fuciformis, and bumblebees, Bombus pasquorum, were observed foraging for nectar in flowers of Viscaria vulgaris. The hawkmoths hovered in front of the flowers, while the bees perched on them. The hawkmoths had a faster probing rate than the bees, and consequently also had higher gross and net rates of energy gain. A model is presented that shows that hovering only yields a higher net rate of energy gain (NREG) than perching when nectar volumes are high due to low competition for the resource. The difference in NREG of perchers and hoverers decreases with an increase of competition, and eventually perching yields the highest NREG. This is an effect of the higher cost of hovering. The results suggest that hovering can only evolve as a pure evolutionarily stable strategy (ESS) if competition is reduced, for example by co-evolutionary specializations with plants. The possibility that it has evolved as a mixed ESS (i.e. individuals can both hover and perch depending on the resource level) is discussed. The evolution of optimal foraging strategies is discussed, and it is pointed out that the rate of gain of an animal is independent of the strategy used when all competing foragers use the same strategy, but competitively superior strategies will nevertheless evolve because they are ESSs. Competition between strategies with different energy costs are special, because resource availability determines which strategy is competitively superior. A high-cost strategy can only evolve as a pure ESS at high resource levels, or as a mixed ESS at intermediate levels.     

Hovering sunbirds in the Old World: occasional behaviour or evolutionary trend?

The nectarivory of sunbirds in the Old World and hummingbirds in the New World evolved independently. While both groups are specialised in their feeding apparatuses, hummingbirds are moreover famous for their adaptations to sustained hovering flight. Recently, an example of a pollination system of the invasive plant Nicotiana glauca has been used to show that less adapted sunbirds also are frequently able to hover. Nevertheless, the question has remained why plants adapted to bird hovering pollination do not occur outside the New World. In this paper we show that the long‐peduncle Cameroonian Impatiens sakeriana is not capable of autonomous selfing and can be pollinated only by two often hovering sunbirds, the Cameroon sunbird Cyanomitra oritis and the northern double‐collared sunbird Cinnyris reichenowi. Our study revealed that this plant is highly specialised for pollination by C. oritis. Cinnyris reichenowi hovers less frequently and often thieves nectar by piercing the flower spur when perching. This study shows that pollination systems occurring in the Old World follow similar evolutionary trends as systems including hovering hummingbirds in the New World.     

Bird-Flowers: Hovering Versus Perching Exploitation

Usually, bird-flowers are classified into New World flowers (used by hovering hummingbirds) and Old World species (adapted to perching birds). Since there are many observations which violate this classification the proposal is made to distinguish between blossoms adapted to hovering birds and those providing a well-defined perch. Characters and examples are given for both groups including the different ways a perch is provided by the plant and the distinct methods of hover-flight in birds. The diverse adaptive solutions for bird pollination within some taxa are exemplified. To understand bird-flowers the actual functioning of flowers must be in focus, and not the geographic distribution nor the systematic affiliation of their visitors.     

Convergent evolution of sunbird pollination systems of Impatiens species in tropical Africa and hummingbird systems of the New World

The bird pollination systems of the New and Old Worlds evolved independently, and differ in many aspects. New World plants are often presented as those adapted to hovering birds while Old World plants to perching birds. Most Neotropical studies also demonstrate that in hummingbird species rich assemblages, only a small number of highly specialized birds exploits the most specialized plants with long corollas. Nevertheless, recent research on bird–plant pollination interactions suggest that sunbird pollination systems in the Old World have converged more with the highly specialized hummingbird pollination systems than previously thought. In this study we focus on the pollination systems of the bird pollination syndrome Impatiens species on Mt. Cameroon, West Africa. We show that despite the high diversity of sunbirds on Mt. Cameroon, only Cyanomitra oritis appear to be important pollinator of all Impatiens species. This asymmetry indicates the absence of pair wise co‐evolution and points to a diffuse co‐evolutionary process resulting in guilds of highly specialized plants and birds; a situation well known from hummingbirds and specialized plant communities of the New World. Additionally, the herbaceous habits of Impatiens species, the frequent adaptations to pollination by hovering birds, and the habitat preference for understory in tropical forests or epiphytic growth, resemble the highly specialized Neotropical plants. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 127–133.     

Dynamic Flight Stability of a Model Hoverfly in Inclined-Stroke-Plane Hovering

Most hovering insects flap their wings in a horizontal plane,called ‘normal hovering'.But some of the best hoverers,e.g.true hoverflies,hover with an inclined stroke plane.In the present paper,the longitudinal dynamic flight stability of a model hoverfly in inclined-stroke-plane hovering was studied.Computational fluid dynamics was used to compute the aerodynamic derivatives and the eigenvalue and eigenvector analysis was used to solve the equations of motion.The primary findings are as follows.(1) For inclined-stroke-plane hovering,the same three natural modes of motion as those for normal hovering were identified:one unstable oscillatory mode,one stable fast subsidence mode,and one stable slow subsidence mode.The unstable oscillatory mode and the fast subsidence mode mainly have horizontal translation and pitch rotation,and the slow subsidence mode mainly has vertical translation.(2) Because of the existence of the unstable oscillatory mode,inclined-stroke-plane hovering flight is not stable.(3) Although there are large differences in stroke plane and body orientations between the inclined-stroke-plane hovering and normal hovering,the relative position between the mean center of pressure and center of mass for these two cases is not very different,resulting in similar stability derivatives,hence similar dynamic stability properties for these two types of hovering.     

The Bizarre Inflorescence of Norantea brasiliensis (Marcgraviaceae): Visits of Hovering and Perching Birds1

The pollination biology of Norantea brasiliensis (Marcgraviaceae) was studied in the rain forest of southeastern Brazil. This plant presents bizarre, brush-type racemous inflorescences bearing numerous flowers and extrafloral cup-shaped nectaries. Flower anthesis is diurnal, nectar production is continuous and copious, and the sticky pollen is readily removed by visitors during the first morning hours. The ruby-coloured inflorescences were visited by eight species of hummingbirds (Trochilidae), and 10 species of passerine birds (three Coerebidae and seven Thraupidae). Hummingbirds hovered while probing for nectar and touched flowers occasionally, whereas passerine birds perched and made contact with flowers habitually. Due to differences in flower-visiting and general foraging behaviour, perching birds act as better pollen vectors than hovering birds. The inflorescence of Norantea brasiliensis seems well fitted for pollination by passerine birds, and the hexose-dominated nectar supports this idea. Pollination syndrome trends within Marcgraviaceae may stem from insect-pollinated, condensed and spike-like inflorescences which would give rise to bird-pollinated, brush-type inflorescences. From the same basic condensed inflorescence, bat-pollinated umbelliform inflorescence may be derived from bird-pollinated, pendulous and corymb-like inflorescences. These postulated inflorescence types are found among the extant species of Marcgraviaceae.     

Interordinal relationships of birds and other reptiles based on whole mitochondrial genomes

Several different groups of birds have been proposed as being the oldest or earliest diverging extant lineage within the avian phylogenetic tree, particularly ratites (Struthioniformes), waterfowl (Anseriformes), and shorebirds (Charadriiformes). Difficulty in resolving this issue stems from several factors, including the relatively rapid radiation of primary (ordinal) bird lineages and the lack of characters from an extant outgroup for birds that is closely related to them by measure of time. To help resolve this question, we have sequenced entire mitochondrial genomes for five birds (a rhea, a duck, a falcon, and two perching birds), one crocodilian, and one turtle. Maximum parsimony and maximum likelihood analyses of these new sequences together with published sequences (18 taxa total) yield the same optimal tree topology, in which a perching bird (Passeriformes) is sister to all the other bird taxa. A basal position for waterfowl among the bird study taxa is rejected by maximum likelihood analyses. However, neither the conventional view, in which ratites (including rhea) are basal to other birds, nor tree topologies with falcon or chicken basal among birds could be rejected in the same manner. In likelihood analyses of a subset of seven birds, alligator, and turtle (9 taxa total), we find that increasing the number of parameters in the model shifts the optimal topology from one with a perching bird basal among birds to the conventional view with ratites diverging basally; moreover, likelihood scores for the two trees are not significantly different. Thus, although our largest set of taxa and characters supports a tree with perching birds diverging basally among birds, the position of this earliest divergence among birds appears unstable. Our analyses indicate a sister relationship between a waterfowl/chicken clade and ratites, relative to perching birds and falcon. We find support for a sister relationship between turtles and a bird/crocodilian clade, and for rejecting both the Haemothermia hypothesis (birds and mammals as sister taxa) and the placement of turtles as basal within the phylogenetic tree for amniote animals.     

Individual differences in reinforcing effects of song

Six out of eight first-year male chaffinches showed some reinforcing effect of song, by perching on a particular perch and remaining on it in order to keep song playing. Since birds could show reinforcing effects by changing frequency, duration, or both, of operant perching, five different indices of reinforcement were considered. Correlations between all but one pair of indices were low, and no index had higher values than any other. For some individuals, one index was high, and for others another. Thus the need for recognizing, rather than ignoring, individual differences is illustrated.     

Auditory prey location in a pause--travel predator: search height, search time, and attack range of Tengmalm's owls (Aegolius funereus)

We studied the nocturnal hunting and diurnal roosting behaviorof 17 radio-equipped Tengmalm's owls (Aegolius funereus), 12males and 5 females, in coniferous forest during their nestingseason. The owls perched lower when hunting than when roosting,probably because hunting perches were selected to minimize thepredator-prey distance or to obtain unobstructed access to theground-dwelling small mammal prey, whereas roosting percheswere selected to minimize the probability of being detectedby an avian predator. There was no difference between perchingheights associated with giving up and prey attack, nor werethere any differences between perching heights, perching times,and attack distances associated with successful and unsuccessfulattacks. There were no sexual differences in perching heightduring hunting or roosting. However, giving-up times tendedto be longer for females than for their mates, which is expectedbecause females are larger than males, and the relative costof flight increases with body mass. The instantaneous attackrate was independent of perching time. The owls gave up theirperches at a constant rate and independently of the amount oftime already spent on the perch in an exponentially decayingpattern. The owls perched longer, however, before launchingan attack than before giving up, probably in order to observedetected prey until the right moment for an attack. Attack distancewas independent of both perching height and perching time. Perchingtime was inversely related to perching height, which fits thetheoretical expectation that the search area will decrease withincreasing height in birds that locate prey auditorily.     

桃红颈天牛成虫的交配行为及对不同寄主植物的栖落和产卵选择性

【目的】桃红颈天牛Aromia bungii是一种钻蛀性害虫,主要以蔷薇科果树和观赏树木为寄主。本研究旨在搞清楚桃红颈天牛成虫的交配产卵行为及其对不同寄主植物的栖落和产卵选择性。【方法】选择苹果Malus pumila、山樱花Prunus serrulata var. spontanea、桃Prunus persica、西府海棠Malus micromalus、毛樱桃Prunus tomentosa、秋子梨Pyrus ussuriensis、杏Prunus armeniaca和红叶李Prunus cerasifera var. atropurea 8种寄主植物木段,构建为成虫的活动场所,观察桃红颈天牛成虫的交配和产卵行为,并测定桃红颈天牛成虫在不同寄主木段上的栖落数量和产卵量。【结果】桃红颈天牛成虫的交配行为分为避让、打斗和交配3种行为。交配过程可分为相遇抱对、插入输精、交配后陪护3个阶段。桃红颈天牛雌虫产卵时无刻槽行为,喜欢将卵产于较为狭窄的缝隙中。雌虫偏好在桃树木段上栖落,雄虫则偏好在毛樱桃木段上栖落。李亚科李属的桃是桃红颈天牛最喜欢的产卵寄主木段,而雌虫在苹果亚科的苹果、海棠和秋子梨木段上产卵极少。相关性分析结果表明雌虫在寄主木段上的栖落数量与产卵量占比间存在正相关性,雄虫在寄主木段上的栖落数量与雌虫在寄主木段上的栖落数量之间也存在正相关性。【结论】桃红颈天牛雌成虫最喜欢桃树木段,雄虫对寄主植物的选择性与雌虫并不完全相同。     

The influence of food on time budgets and timing of breeding of the Dunnock Prunella modularis

In two years, we provided some female Dunnocks with extra food from January through to July. In one year fed females bred ten days earlier than controls and in another year they bred 22 days earlier, but in neither year did they lay larger clutches. Matched comparisons of the same females on the same territories, who had food in one year but not in the other, showed the same effects. Within both feeder and control females, the earliest breeders were those which had spent more time perching in late winter. Perching time may be a good measure of the time an individual has available above that needed for self maintenance. Therefore females who spent more time perching may have been those first able to cross the threshold of extra time needed for the start of breeding activities.     

Lipid economy,flight activity and reproductive behaviour in the speckled wood butterfly: on the energetic cost of territory holding

Insect flight is a highly energy demanding type of locomotion. In butterflies, males may locate females by different behavioural tactics. The tactics correspond to different flight types that, in turn, are assumed to reflect different energetic costs. Costs need to be considered to fully understand the pay‐offs of co‐existing alternative tactics relative to the environmental context and the phenotypes of the individuals. We addressed the issue in the speckled wood Pararge aegeria, in which males either adopt a territorial wait‐and‐fight tactic (i.e. territorial perching) in a sunlit patch on the forest floor, or a fly‐and‐search tactic to locate females in a wider area of the forest (i.e. patrolling). Perching corresponds to high frequency of take‐off flights and aerial combats with high levels of manoeuvrability and is assumed to be energetically more costly than longer, continuous flights at lower speed in patrollers. We tested the effect of different flight activity levels and of the behavioural tactics on lipid reserves and lipid use in males by laboratory and outdoor cage experiments. Low‐activity males that had access to honey water were capable of synthesizing lipids; their lipid reserves increased with age. The effect disappeared in males that actively flew in the outdoor cages. Lipid reserves decreased significantly faster in territorial perching males compared to non‐perching males, but resting metabolic rate did not differ between the alternative behavioural tactics. Territorial perching males had larger flight muscle ratio (i.e. thorax/body mass) than non‐perching males. We discuss the evidence of the physiological costs of perching relative to the co‐existence of perching and patrolling tactics.     

SEXUAL SELECTION AND BUTTERFLY DESIGN—A COMPARATIVE STUDY

Temperate butterflies of 44 species were examined to determine if their mating system (perching and patrolling) affected flight design. To control for spurious effects due to ancestry, 25 of these species were assigned to eight contrasts within which a change in mating system had occurred. In perching species sexual selection was predicted to favor traits associated with high acceleration ability and speed, while in patrolling species traits associated with flight endurance were predicted. In conformance with these expectations males of perching species had larger thorax/body mass ratios, higher wing loadings, and higher aspect ratios than patrolling species. The male mating system affected females in the same direction in the same variables as males. This could be explained by a genetic correlation with males. When removing the covariance between the sexes, only male design was explained by the mating system. The mating system was also associated with different degrees of sexual dimorphism in wing size. This supported the hypothesis that male design was affected by the mating system.     

Introducing a new tool to derive animal behaviour from GPS data without ancillary data: The red-footed falcon in Italy as a case study

Animal monitoring is a key stage in the process of understanding individual behaviour and social dynamics that characterize populations and species. Recent technological advances allow to collect large data sets on animal movement. Using such data to compute activity patterns is very challenging without expensive field surveys, video analyses and/or accelerometer sensors.We introduce here a new tool (UABE: Unsupervised Animal Behaviour Examiner) that makes use of a set of methodologies (if-then-else rules, sensitivity analyses, Markov chains) to interpret GPS data in behavioural terms under worst case conditions, i.e. in absence of ancillary data. The UABE is halfway between unsupervised inductive (clustering or segmentation) and deductive (expert-based) methods. As a case study, we applied the UABE to the investigation of the red-footed falcons' activity at nest at the two largest breeding colonies in Italy during the chick-rearing period.By using accurate GPS data-loggers, we tracked four adults in June–July 2019 and 2020, and collected 5840 GPS points. Activity at nest occupied one fourth of the red-footed falcons' day, with substantial differences between months and between daytime and night-time. On an hourly basis, clear temporal patterns of activity at nest emerged. Through the combined use of frequency and continuity of the studied behaviour, the UABE also allowed to raise plausible hypotheses for the reasons behind birds' activity at nest at specific times of day.The UABE supports the creation of unsupervised ethograms of both stationary (e.g., standing at nest or roost, hovering/perching) and nonstationary (e.g., flight, walking) activity patterns. It can be applied to any species regardless of the tracking instrument used.     

Perching of Tengmalm's Owl (Aegolius funereus) Nestlings at the Nest Box Entrance: Effect of Time of the Day,Age, Wing Length and Body Weight

The behaviour of the nestlings of nocturnal cavity-nesting species has relatively rarely been studied in detail because of problems connected with use of the technical devices required to provide long-term monitoring of individuals. However, long-term observation of nestling behaviour is crucial in order to identify different types of behaviour which may be caused by sibling competition at the end of nesting period. We studied behaviour of 43 Tengmalm''s owl (Aegolius funereus) nestlings at 14 nests using a camera and a chip system. The nestlings perched at the nest box entrance from an average age of 28 days from hatching (range 24–34 days) until fledging, spending around 2 hours per day here in total, in periods ranging from a few seconds to 147 min (7.6±10.9 min, mean ± SD). We found that individual duration of perching at the nest box entrance was significantly influenced by nestlings'' age and wing length and that the duration of perching at the nest box entrance significantly decreased with time of night. However, during daylight hours, time of day had no effect on either probability or duration of nestlings'' perching. We suggest daylight perching at the nest box entrance results from nestlings'' preparation for fledging, while individuals perching here during the night may gain an advantageous position for obtaining food from the parents; another possibility at all times of day is that nestlings can reaffirm their social dominance status by monopolizing the nest box entrance.     

Timing of mate-locating by males in relation to female activity in the carpenter beeXylocopa varipuncta (Hymenoptera: Apidae)

Males of the carpenter beeXylocopa varipuncta Patton wait for females to visit them as they hover at landmark territories along ridgelines on some spring afternoons. While hovering, males advertize their presence by releasing a pheromone that attracts passing females. If males have limited time to invest in territorial hovering and signaling, then they should engage in these activities more often at times when mate-searching females are most likely to visit landmark territories. The number of females flying near male territories varies greatly over the course of afternoons and from day to day. Measures of female activity and male territorial activity were highly correlated at one study site, both in terms of changes within afternoons and in terms of day-to-day fluctuations. This result supports the hypothesis that males ofX. varipuncta time their mateattracting behavior to maximize contacts with receptive females.     

Perturbation analysis of 6DoF flight dynamics and passive dynamic stability of hovering fruit fly Drosophila melanogaster

Insects exhibit exquisite control of their flapping flight, capable of performing precise stability and steering maneuverability. Here we develop an integrated computational model to investigate flight dynamics of insect hovering based on coupling the equations of 6 degree of freedom (6DoF) motion with the Navier-Stokes (NS) equations. Unsteady aerodynamics is resolved by using a biology-inspired dynamic flight simulator that integrates models of realistic wing-body morphology and kinematics, and a NS solver. We further develop a dynamic model to solve the rigid body equations of 6DoF motion by using a 4th-order Runge-Kutta method. In this model, instantaneous forces and moments based on the NS-solutions are represented in terms of Fourier series. With this model, we perform a systematic simulation-based analysis on the passive dynamic stability of a hovering fruit fly, Drosophila melanogaster, with a specific focus on responses of state variables to six one-directional perturbation conditions during latency period. Our results reveal that the flight dynamics of fruit fly hovering does not have a straightforward dynamic stability in a conventional sense that perturbations damp out in a manner of monotonous convergence. However, it is found to exist a transient interval containing an initial converging response observed for all the six perturbation variables and a terminal instability that at least one state variable subsequently tends to diverge after several wing beat cycles. Furthermore, our results illustrate that a fruit fly does have sufficient time to apply some active mediation to sustain a steady hovering before losing body attitudes.