Honeyeater foraging: A test of optimal foraging theory

Honeyeaters (Meliphagidae) were observed foraging for nectar from Lambertia formosa inflorescences, each of which has seven flowers. The frequency distribution of numbers of flowers probed per visit to an inflorescence was found to be bimodal, with one peak at two and the other at seven. It is hypothesized that this frequency distribution results from a rule of departure from inflorescences that maximizes the net rate of energy gain. Patterns of nectar distribution were determined for a large sample of inflorescences. In addition the extent to which the honeyeaters re-probe flowers during a visit to an inflorescence was estimated. From these data and from field measurements of the times required by the honeyeaters to perform the various foraging behaviours, computer simulations of honeyeater foraging were constructed. These simulations led in turn to optimal frequency distributions of numbers of flowers probed per inflorescence that were bimodal but had peaks at 1 and 7 instead of 2 and 7. Although the observed and predicted behaviour were consequently similar, the difference between them was nevertheless significant. This difference could have been due to the birds' transient occupancy of the study area.     

Optimal Foraging in Hummingbirds: Testing the Marginal Value Theorem

To a hummingbird, clusters of flowers on inflorescences representpatches and provide an ideal situation to test prediction ofoptimal patch-use. The basic question is what decision ruleshould a hummingbird use to decide whether or not to leave aninflorescence? The hypothesis is that hummingbirds will adoptthe decision rule that maximizes their net rate of energy gainwhile foraging. This hypothesis leads to an analogue of Charnov'smarginal value theorem which determines an optimal decisionrule. The optimal decision ruleis then used to predict aspectsof the hummingbirds' foraging, and these predictions are comparedwith field data The optimal decision rule is a function of how much informationis used by the hummingbirds. Data indicate that a decision toleave an inflorescence is a function of the number of flowersvisited, the number of flowers available on the inflorescence,and the amount of nectar obtained at the last flower. The optimaldecision rule was calculated assuming no additional informationis used.     

Pollen foraging by bumblebees: Foraging patterns and efficiency on Lupinus polyphyllus

Summary Bumblebees foraging on vertical inflorescences start near the bottom and work upward, behavior commonly interpreted as a response to the greater amounts of nectar available in lower flowers. Lupinus polyphyllus, which produces no nectar, has more pollen available in upper flowers. Although bees are probably unable to detect this gradient, since pollen is hidden from their view, they still start low and forage upward. Therefore, we concluded that the bees' tendency to forage upward on vertical inflorescences is not tied to a reward gradient. In addition, bees use only about 15% of the flowers per inflorescence, although they could be much more efficient by visiting and revisiting every flower systematically. In general, revisits would not be penalized because most flowers contain enough pollen for several visits. Optimal foraging theory may not offer an adequate explanation for such gross inefficiency.     

珍贵绢丝昆虫——天蚕蚕蛾翅振行为观察

本文较系统地观察和测定了天蚕蚕蛾的翅振时间节律和翅振频率。结果表明,雄蚕蛾不论独处还是与雌蚕蛾共存,其翅振行为均出现于18：00至次日6：00之间,其中以22：00至次日4：00为主;雌蚕蛾独处时未见翅振行为,只有与雄蛾共存时方有此行为,出现于22：00至次日6：00。雌雄蚕蛾翅振行为参数,包括日翅振次数、翅振总时间、一次翅振持续时间、两次翅振间间隔时间等,因蚕蛾日龄和存在方式(独处或与异性共存)而异;雌雄蚕蛾相比,除两次翅振间隔时间外,雄蛾均明显高些。雌雄蚕蛾翅振频率一般为7-12Hz,不同日龄闯,雄蛾以羽化第2天为高,平均10.9741Hz;雌蛾以羽化第1天为高,平均10．7071Hz。同一日龄不同时刻间翅振频率有一定差异,但未达显著水平。此外,尚观察了雄蛾触角对其翅振行为表现的作用,最后讨论了制种中拟注意的有关问题。     

Simulating avian wingbeat kinematics

Inverse dynamics methods are used to simulate avian wingbeats in varying flight conditions. A geometrically scalable multi-segment bird model is constructed, and optimisation techniques are employed to determine segment motions that generate desired aerodynamic force coefficients with minimal mechanical power output. The results show that wingbeat kinematics vary gradually with changes in cruise speed, which is consistent with experimental data. Optimised solutions for cruising flight of the pigeon suggest that upstroke wing retraction is used as a method of saving energy. Analysis of the aerodynamic force coefficient variation in high and low speed cruise leads to the proposal that a suitable gait metric should include both thrust and lift generation during each half-stroke.     

Morphological correlates of nectar production used by honeybees

Abstract.  1. Honeybees foraging on lavender have been shown to choose inflorescences that are larger and have more flowers. If they are selecting optimally then these inflorescences should yield higher net rates of energy gain. The number and distribution of flowers within inflorescences is a complex function of age, however, which might itself influence nectar quality and availability.
2. Sampling of the overnight nectar secretion of visited and unvisited inflorescences showed that younger inflorescences with more flowers produced more sugar per flower and had fewer unproductive flowers than other inflorescences, but the size of the inflorescence had no effects.
3. Overall display size attracted bees to inspect inflorescences, as inflorescences that were inspected but rejected were larger and/or had larger or more bracts than those that were ignored. Bees, however, accepted more productive inflorescences based on different cues: inflorescence age and number of flowers.
4. Inflorescence choice thus appeared to reflect a two-stage decision process based on different morphological criteria at each stage.     

Geitonogamy in rewarding and unrewarding inflorescences: modelling pollen transfer on actual foraging sequences

Many orchid species are unusual in that they provide no nectar or pollen rewards for their pollinators. Absence of reward is expected to have a fundamental effect on pollinator visitation patterns. In particular the number of flowers visited per inflorescence is expected to be affected in both unrewarding and co-flowering rewarding species. We used arrays of artificial inflorescences, which could be either rewarding or unrewarding and were differentiated by their colour, to test how many flowers bumblebees visit in each type of inflorescence. The frequency of the two colours was varied, thus modelling the case where different frequencies of both an unrewarding and rewarding species were present in a patch. We found that bumblebees visited more flowers per rewarding inflorescence after they have experienced unrewarding or partially emptied rewarding inflorescences. We used these results to simulate pollen transfer and thus predict selfing rates on rewarding inflorescences. We found these increased when nectar depleted or when there was a greater proportion of unrewarding inflorescences in the patch. Conversely, we found that the number of flowers bumblebees visited on each unrewarding inflorescence did not significantly change through experiments. Selfing rates for unrewarding inflorescences were predicted to depend principally on the number of these inflorescences bumblebees visited rather than on the number of flowers they visit per inflorescence. This was because most visitors to orchids are supposed to be naive, and pollinators that commence foraging carrying no pollen will necessarily self any flower they pollinate on the first inflorescence they visit. Thus the average selfing rate is expected to increase as the sequence of inflorescences visited decreases in length.     

Optimal foraging in hummingbirds: Rule of movement between inflorescences

The movements of hummingbirds between inflorescences of scarlet gilia (Ipomopsis aggregata) were studied. These movements exhibited the following patterns: (1) Although the hummingbirds appeared to avoid moving to the previous inflorescence, no significant correlation was found between the directions of successive inter-inflorescence movements. (2) The frequency distribution of inter-inflorescence flight distances was found to be leptokurtic. (3) The hummingbirds were more likely to move to an inflorescence the larger and/or closer it was. (4) The hummingbirds moved to inflorescences of greatest apparent size (i.e. ratio of number of flowers available to distance from present inflorescence) more often than they moved to the largest inflorescence, the closest infloresence, or the inflorescence estimated to yield the greatest rate of energy gain. (5) The frequency distribution of moves to the inflorescence having the ith greatest apparent size is well fitted by a geometric distribution. This is consistent with the hummingbrids choosing the inflorescence of greatest apparent size (excluding the previous inflorescence) from within some scanning sector. These movement patterns are consistent with the expectations of optimal foraging theory only if the hummingbirds cannot or do not determine the directions of possible inflorescences relative to the direction of arrival at the present inflorescence and if they cannot assess independently the sizes and distances of possible inflorescences.     

Accumulating wing damage affects foraging decisions in honeybees (Apis mellifera L.)

Abstract.  1. Nectar-foraging honeybees ( Apis mellifera ) on lavender ( Lavandula stoechas ) appear to forage so as to maximise net energy return from foraging bouts; however, evidence from other studies suggests that foraging has a detrimental effect on survival, due at least in part to physiological deterioration of the flight mechanism. But foragers also acquire wing damage during foraging, which may increase foraging effort and reduce foraging lifespan.
2. The accumulation of damage over time and its effects on foraging flight and flower choice were studied in the field using a system in which the criteria for flower preference by foragers was known from previous work. Wing damage accumulated exponentially over time and resulted in foragers becoming less choosy about the flowers they visited.
3. Damage added experimentally contributed independently to the effect on choosiness. Effects of wing damage (natural and added experimentally) were also independent of those of a relative measure of age, which related in an inconsistent way to changes in foraging preferences.     

黄猄蚁对大蜜蜂的捕食行为及其潜在影响

捕食作用会对访花昆虫的种群、行为以及植物适合度产生影响,是植物与传粉者相互关系研究中常被忽视的因素.本文报道了黄猄蚁对大蜜蜂的捕食行为,并模拟捕食的关键环节研究了捕食过程对重要访花昆虫行为的影响.结果表明,黄猄蚁能够利用局部环境主动攻击猎物,利用群体合作捕获采集过程中的体型较大的大蜜蜂,捕食威胁是其影响植物-访花者关系的重要机制.大蜜蜂具有感知花上危险的能力,模拟处理的个体会逃离危险的花或植株并在花上留下标记,将危险信息传递给其它个体.其它拜访者对具有危险信号花的采集频次明显减少,采集时间缩短;模拟处理的影响会随时间推移而较快地消失.此外,该实验没有发现大蜜蜂在花上停留采集过程中具有明显的防御行为.     

Coordination of wingbeat and respiration in birds. II. "Fictive" flight.

To determine whether an interaction between central respiratory and locomotor networks may be involved in the observed coordination of wingbeat and respiratory rhythms during free flight in birds, we examined the relationship between wingbeat and respiratory activity in decerebrate Canada geese and Pekin ducks before and after paralysis. Locomotor activity was induced through electrical stimulation of brain stem locomotor regions. Respiratory frequency (fv) was monitored via pneumotachography and intercostal electromyogram recordings before paralysis and via intercostal and cranial nerve IX electroneurogram recordings after paralysis. Wingbeat frequency (fW) was monitored using pectoralis major electromyogram recordings before, and electroneurogram recordings after, paralysis. Respiratory and cardiovascular responses of decerebrate birds during active (nonparalyzed) and "fictive" (paralyzed) wing activity were qualitatively similar to those of a variety of vertebrate species to exercise. As seen during free flight, wingbeat and respiratory rhythms were always coordinated during electrically induced wing activity. Before paralysis during active wing flapping, coupling ratios (fW/fv) of 1:1, 2:1, 3:1, and 4:1 (wingbeats per breath) were observed. After paralysis, fW and fv remained coupled; however, 1:1 coordination predominated. All animals tested (n = 9) showed 1:1 coordination. Two animals also showed brief periods of 2:1 coupling. It is clear that locomotor and respiratory networks interact on a central level to produce a synchronized output. The observation that the coordination between fW and fv differs in paralyzed and nonparalyzed birds suggests that peripheral feedback is involved in the modulation of a centrally derived coordination.     

Optimal foraging in bumblebees and coevolution with their plants

Summary The aims of this paper were to consider the coevolution between bumblebee movement patterns within plants and various properties of the plants such as the spatial distribution of their flowers, and to determine the extent to which the bumblebees and the plants can be considered to be maximally adaptive or optimal. Attention was restricted to plants which have flowers arranged on vertical inflorescences and to the bumblebees which visit these plants.It was found that the bumblebees tend to commence foraging at the bottom of each infloresence, that they tend to move from one flower to the closest vertically higher flower, that they miss flowers as they move upwards and that they tend to leave each inflorescence before reaching the top. It was also found for the four common plant species considered that nectar abundance per flower decreases with flower height on an inflorescence, that the flowers with receptive stigmas are restricted to the bottoms of the inflorescences while the flowers shedding pollen occur above them, and that the flowers are arranged approximately in spirals on the inflorescences.The pattern of movements of the bumblebees and the various properties of the plants appear to represent coevolved adaptations. Furthermore the bumblebees' movement patterns appear to be optimal in the sense that they result in the maximum net rate of energy gain to the bumblebees. Further studies are necessary, however, to determine whether or not the plants can be considered to be optimal.An exception to the above scheme is provided by a plant which is quite uncommon in the study area. This plant also has flowers on vertical inflorescences and appears to be pollinated by bumblebees. However, while the pattern of movements of the bumblebees on this plant species are extremely similar to those on the four common species, this plant species exhibits quite different properties from the other four. Two possible explanations for this exception are presented.     

铁破锣花序的开花特性与昆虫传粉的相互关系研究

为了揭示植物花的空间布局与开花动态的调节机制以及避免同株异花传粉的生态学策略,该研究对铁破锣［Beesia calthifolia (Maxim.) Ulbr.］花序形态结构、开花动态和传粉生物学进行了观察分析。结果表明：（1）铁破锣花序结构设计巧妙,由3朵花组成一个聚伞花序单元并依次排列在主花序轴上,且花序轴上聚伞花序之间距离较远。（2）铁破锣通过单个聚伞花序顶花先开,通常只有6～8朵聚伞花序的顶花同时开放,而且总状花序从基部到顶部逐次开放,从而使得大量聚集单花的花序达到尽量少开花。（3）铁破锣花白色,花粉是访花昆虫的仅有诱物,纤细巴蚜蝇（Baccha maculata）是铁破锣的主要传粉昆虫,这种昆虫能够以花丝为着力点取食花粉,通常在一个花序上取食一朵单花后很快飞向另外一个花序的花。研究认为,铁破锣花序的空间设计和开花的时间序列动态减少了昆虫访问同株异花的可能性。     

Wingbeat frequency and flap-pause ratio during natural migratory flight in thrushes

Powered flapping flight has evolved independently in many differenttaxa. For flapping fliers, wingbeat parameters such as frequencyand amplitude are the primary determinants of these animals’energetic expenditure during flight. Here we present data onwingbeat frequency and amplitude for three New World thrushspecies during 15 entire nocturnal migratory flights over theMidwestern United States. Using continuous (non-pulsing) radiotransmitters, we were able to measure wingbeat frequency andrelative amplitude of wingbeats as well as the characteristicsof flap-pauses. Contrary to previous telemetric findings, allof the individuals we followed used both flapping-only and flap-pauseflight. During migratory flights, wingbeat frequency, effectivewingbeat frequency, and amplitude were highest during initialascent. Effective wingbeat frequency and amplitude were lowestduring final descent. We show that identification of speciesbased solely on characteristics of the wingbeat e.g., duringradar studies, can be difficult because variables such as wingbeatfrequency and amplitude, wingbeat pausing, and pattern of beatsand pauses vary between individuals of the same species andeven within individual flights. We also show that observed wingbeatfrequencies were lower than those predicted by theoretical models.We speculate that this may be because theoretical predictionsare generally based on (1) data from larger birds and (2) datafrom diurnal flights. We found that diurnal wingbeat frequenciesof thrushes were generally higher than were those during nocturnalmigratory flight. Finally, we suggest that rather than remainingat a single altitude during flight or climbing slightly as theoreticalmodels predict, thrushes often moved up and down in the aircolumn, perhaps searching for favorable atmospheric conditions.     

Manipulation of bee behavior by inflorescence architecture and its consequences for plant mating

Angiosperms display flowers in many three-dimensional arrangements, but the functional significance of this diversity is largely unknown. We examined influences of inflorescence architecture on pollination and mating by quantifying the responses of bumblebees to three architectures and then using these observations as the basis of a model that simulated pollen dispersal. On artificial panicles, racemes, and umbels, each with 12 identical flowers, bees visited one more flower, on average, on umbels than on panicles (with racemes being intermediate). In contrast to this weak response, the consistency of foraging paths among flowers differed strongly among architectures (raceme > panicle > umbel). The simulation model revealed limited differences in self-pollination and pollen export among the three inflorescence designs when all flowers presented and received pollen, because mating differences depended on only the number of flowers visited. In contrast, in simulations of inflorescences on which pollen receipt and presentation were segregated so as to minimize interference among flowers, the consistency of movement paths governed mating. In this case, racemes self-pollinated much less than umbels (with panicles being intermediate), and racemes exported much more pollen than umbels and panicles. These effects have diverse consequences for the evolution of inflorescence architecture, flower design, and sexual segregation.     

The flight performance of tsetse flies.

The four species of tsetse used are shown to be aerodynamically rather similar. The wingbeat frequency, aerodynamic lift, flight duration, and number of wingbeats per flight are seen to undergo large increases during adult maturation. Tsetse can produce unusually large aerodynamic lifts, although the flight performance declines very quickly during individual flights. The large lifts appear to be associated with the high percentage of muscle in the fly, and the rapid decline with the inability of the fly to replenish rapidly its proline reserves. Wing mutilation has been shown to increase the wingbeat frequency. The potentiation of the flight performance during maturation appears to be causally linked to increases in the thoracic muscle, cuticle, and proline levels. The results presented here are at variance with those of an earlier study on the physics of flight in tsetse.     

The movement patterns of carpenter beesXylocopa micans and bumblebeesBombus pennsylvanicus onPontederia cordata inflorescences

The movement patterns of carpenter bees (Xylocopa micans) and bumblebees (Bombus pennsylvanicus) foraging for nectar on vertical inflorescences ofPontederia cordata were studied near Miami, Florida. The floral biology ofP. cordata is unique in several ways: (a) many short-lived flowers per inflorescence, (b) constant nectar production throughout the life span of each flower, and (c) abscence of vertical patterning of nectar and age of flowers. Inflorescences ranged between 3.5 and 15.8 cm long and had between 9 and 55 open flowers. Both carpenter bees and bumblebees arrived mostly on the bottom third of the inflorescence and left after visiting flowers on the top third of the inflorescence. The departure position from the inflorescence was higher up than observed in studies of other insect pollinators foraging on other speces of plants. This pattern of departure probably occurs in the absence of a vertical gradient of nectar or floral morphology.     

Caste-specific patterns of flower visitation in bumble bees (Bombus kirbyellus) collecting nectar from Polemonium viscosum

ABSTRACT.

• 1 Foraging routes of worker and queen bumble bees (Bombus kirbyellus Curtis) collecting nectar from flowers of the alpine sky pilot, Polemonium viscosum Nutt., were followed and the corolla tube length, corolla diameter, floral scent, and number of flowers on plants visited or bypassed by bees were monitored. Additionally, the number and proportion of flowers visited per inflorescence and distance flown from each to the next were recorded. Queens and workers differed significantly in choice of flowers. However, intra-inflorescence visitation rates and departure distances were similar between castes. Castes differed in the extent to which visitation reflected patch quality versus individual floral traits.
• 2 Both queens and workers failed to visit skunky-flowered plants more often than they failed to visit sweet-flowered ones, and preferred large over small inflorescences. However, queens visited large-flowered plants more often than small-flowered ones, while workers preferred flowers with shorter corolla tubes, regardless of their diameter. Although a number of studies have documented caste specialization on alternate species of host plants, ours is one of the first to show that morphological preferences promote comparable foraging differences between castes on monospecific plant resources.
• 3 Queens, once on a plant, responded to floral traits by probing more flowers on large inflorescences, as well as on those with broader floral form. Workers did not alter intra-inflorescence visitation rate in response to floral traits.
• 4 For workers, no significant relationship was demonstrated between the likelihood of passing by a plant and the number of flowers probed on the previous inflorescence visited. Thus, workers appeared to accept or reject each plant of P. viscosum independently. However, queens passed by fewer plants when leaving rich inflorescences than poor ones. These results suggest that workers use only individual plant acceptability in choosing which plants to visit, whereas queens base plant choice on patch and individual attributes. Such differences between castes in foraging rules when exploiting the same floral resource have received little attention, and provide insights into the heterogeneity of harvestable reward distributions from the perspective of the forager population.

     

金斑蝶觅食求偶过程中的视觉嗅觉利用

【目的】阐明金斑蝶Danaus chrysippus觅食求偶时视觉嗅觉信息利用机制。【方法】通过行为学观察,测试金斑蝶成虫对7种不同颜色的假花和假花喷施10%蜂蜜水后的觅食响应;通过自然种群求偶和塑封模型访问试验,验证金斑蝶对颜色和气味行为响应。【结果】在无气味的7种颜色的假花条件下,金斑蝶成虫访花总次数为138次,对长波长红色(620～750 nm)显示出明显的颜色趋性(占总访花次数的 52.17%),访问蓝色(占总访花次数的15.22%)和橙色(占总访花次数的11.59%)次之,其他4种颜色假花较少访问;在假花上喷洒10%蜂蜜水后,金斑蝶成虫访花总次数增加3.05倍,雌雄蝶访花次数分别增加3.31和2.86倍;去花冠无气味的花枝几乎没有蝴蝶访问,然而在去花冠无气味的假花花枝上喷洒10%蜂蜜水后,金斑蝶成虫访花总次数增加146次,对单一颜色的访问次数与对单一气味的访问次数相比无显著差异,金斑蝶成虫对气味加颜色的访问总次数较对单一气味的访问总次数增加2.88倍,雌雄蝶访花次数分别增加2.95和2.83倍;金斑蝶雄蝶对颜色和气味的敏感性高于雌蝶。在求偶过程中,金斑蝶对自然种群与蝶翅模型都有雄追雄、雄追雌、雌追雌和雌追雄4种模式,但金斑蝶对无气味蝶翅模型的追逐次数显著低于对自然种群的追逐次数。在自然光下,雌雄蝶翅颜色和图案相近,除了雄蝶后翅腹面多一白色性标斑点外,其他无显著区别;在紫外光下,蝶翅模型的颜色和图案比自然蝶翅更为明显,可能有助于异性识别。【结论】金斑蝶成虫在觅食和求偶过程中,视觉嗅觉同等重要。颜色较稳定、醒目、容易被发现;气味不稳定、易挥发、受环境干扰较大。在觅食中,金斑蝶利用视觉先进行初步定位,通过嗅觉和味觉确认食物资源;在求偶中,金斑蝶成虫通过视觉初步区分异性,然后依靠嗅觉进行精确识别。     

Flower choice by honey bees (Apis mellifera L.): sex-phase of flowers and preferences among nectar and pollen foragers

Bees foraging for nectar should choose different inflorescences from those foraging for both pollen and nectar, if inflorescences consist of differing proportions of male and female flowers, particularly if the sex phases of the flowers differ in nectar content as well as the occurrence of pollen. This study tested this prediction using worker honey bees (Apis mellifera L.) foraging on inflorescences of Lavandula stoechas. Female flowers contained about twice the volume of nectar of male flowers. As one would predict, bees foraging for nectar only chose inflorescences with disproportionately more female flowers: time spent on the inflorescence was correlated with the number of female flowers, but not with the number of male flowers. Inflorescence size was inversely correlated with the number of female flowers, and could be used as a morphological cue by these bees. Also as predicted, workers foraging for both pollen and nectar chose inflorescences with relatively greater numbers of both male and female flowers: time spent on these inflorescences was correlated with the number of male flowers, but not with the number of females flowers. A morphological cue inversely associated with such inflorescences is the size of the bract display. Choice of flowers within inflorescences was also influenced predictably, but preferences appeared to be based upon corolla size rather than directly on sex phase.