Trapline foraging by bumble bees: III. Temporal patterns of visitation and foraging success at single plants

We analyzed the temporal structure of visitation by bumble beeworkers to a single Penstemon strictus plant growing in an arrayof conspecifics. When tested against a null distribution usinga randomization model, the observed pattern of arrivals forthe whole group of bees was random, but departures were clusteredin time. Certain individuals visited the plant repeatedly andfrequently throughout the day. These showed significantly regulararrival and departure schedules, which were likely producedby traplining. We explored whether these more frequent and regularforagers gained a higher reward than random or incidental plantvisitors. Using an analytical model, Possingham predicted thata dominating forager that visited a simple, renewing resourcein a regular pattern would garner higher and less variable rewardsthan random visitors. Inspired by these results, but interestedin plant-level visitation, we constructed a simulation modelof resource dynamics for a multiflowered plant with high visitation.The model incorporates the observed visitation schedules ofall bees and independent reward dynamics for each flower onthe plant. We calculated the rewards that observed bees wouldhave collected given a range of resource-renewal parameters.More frequent visitors did not return to the plant when whole-plantresource levels were higher, but these visitors did get greaterrewards. Their increased reward resulted from greater foragingefficiency, primarily through selecting (on average) more rewardingflowers than those selected by less frequent, random visitors.     

Trapline foraging by bumble bees: IV. Optimization of route geometry in the absence of competition

Foraging on resources that are fixed in space but that replenishover time, such as floral nectar and pollen, presents animalswith the problem of selecting a foraging route. What can flowervisitors such as bees do to optimize their foraging routes,that is, reduce return time or route distance? Some repeatedlyvisit a set of plants in a significantly predictable sequence(so-called "trapline foraging"), which may also enhance theirforaging efficiency. A moderate level of optimization and repetitionof foraging routes can be reached by following simple movementrules for choosing the distances and turning angles of successiveflights, without the use of spatial memory. If pollinators canlearn the locations of patches and choose among possible foragingroutes or paths, however, even better performance may be achieved.We tested whether and how bumble bees can optimize and repeattheir foraging routes in laboratory experiments with artificialflowers that secreted nectar at a constant rate. With increasingexperience, foraging routes of bees became more repeatable andefficient than expected from a combination of simple movementrules between successive flowers. We suggest that trapline foragingis a more sophisticated pattern of spatial use than searchingand is based on memory. On the other hand, certain spatial configurationsof flowers hampered optimization by the bees; bees preferredto choose short distances over straight moves and showed littleplasticity in this regard. Developing an efficient trapline,therefore, may require prior selection of a set of plants withan appropriate spatial configuration.     

Trapline foraging by bumblebees: I. Persistence of flight-path geometry

By setting out arrays of potted plants of Penstemon strictus,I tested whether freely foraging bumblebee (Bombus spp. ) workerswould establish regular foraging routes that reflected the geometryof the array. They did, passing through an asymmetrical arrayin a pattern that minimized interplant flight distances. Afterthe array was changed to a symmetrical pattern, however, theexperienced bees continued to show their previous asymmetricalflight patterns. New bees without experience on the asymmetricalarray showed no asymmetry on the symmetrical array. I term thispersistence of flight-path geometry "trapline holdover, " anddiscuss its implications for the study of animals' learningand foraging behavior.     

Bees do not use nearest-neighbour rules for optimization of multi-location routes

Animals collecting patchily distributed resources are faced with complex multi-location routing problems. Rather than comparing all possible routes, they often find reasonably short solutions by simply moving to the nearest unvisited resources when foraging. Here, we report the travel optimization performance of bumble-bees (Bombus terrestris) foraging in a flight cage containing six artificial flowers arranged such that movements between nearest-neighbour locations would lead to a long suboptimal route. After extensive training (80 foraging bouts and at least 640 flower visits), bees reduced their flight distances and prioritized shortest possible routes, while almost never following nearest-neighbour solutions. We discuss possible strategies used during the establishment of stable multi-location routes (or traplines), and how these could allow bees and other animals to solve complex routing problems through experience, without necessarily requiring a sophisticated cognitive representation of space.     

The trouble with isolation by distance

The genetic population structure of many species is characterised by a pattern of isolation by distance (IBD): due to limited dispersal, individuals that are geographically close tend to be genetically more similar than individuals that are far apart. Despite the ubiquity of IBD in nature, many commonly used statistical tests are based on a null model that is completely non-spatial, the Island model. Here, I argue that patterns of spatial autocorrelation deriving from IBD present a problem for such tests as it can severely bias their outcome. I use simulated data to illustrate this problem for two widely used types of tests: tests of hierarchical population structure and the detection of loci under selection. My results show that for both types of tests the presence of IBD can indeed lead to a large number of false positives. I therefore argue that all analyses in a study should take the spatial dependence in the data into account, unless it can be shown that there is no spatial autocorrelation in the allele frequency distribution that is under investigation. Thus, it is urgent to develop additional statistical approaches that are based on a spatially explicit null model instead of the non-spatial Island model.     

Response of a traplining hummingbird to changes in nectar availability

Theory predicts that nectarivores respond to changes in profitabilityof patches of flowers or feeders by adjusting visitation rateto increase reward size. We conducted a set of experiments inan outdoor enclosure with seven feeders to determine how Phaethornislongirostris, a traplining hummingbird, adjusts its visitationrates in response to changes in sucrose solution delivery rates.Each experiment tested the response of P. longirostris to thefollowing changes in the timing and volume of sucrose solutiondelivery: (1) increases in sucrose solution abundance at allfeeders (mimicking seasonal increases in numbers of open flowersor nectar output); (2) large changes in sucrose solution availabilityat one feeder (mimicking increases or decreases of patch profitability);and (3) sudden unexpected decreases in sucrose solution availabilityat one feeder (mimicking loss of nectar to competitors). Wefound that P. longirostris (1) decreased visitation rates whenthe sucrose solution delivery rate was higher at all feeders,(2) increased visitation rates to individual feeders when their profitabilityincreased for whole days but did not significantly decrease visitationrates when feeder output decreased; and (3) responded to sudden foodlosses at a feeder (due to simulated competition) by increasinguse of that feeder for 1 or 2 h after the loss.     

A comparison of bumblebees' movements in uniform and aggregated distributions of their forage plant

1. Movements by bumblebees Bombus lapidarius were studied in arrays of plants of oil‐seed rape Brassica napus cv Westar. In the arrays, plants were arranged into either a uniformly dispersed configuration or a patchily dispersed configuration. 2. Progress across an array was measured as the distance bumblebees had moved away from an original plant location after eight subsequent, successive plant visits. Movements were analysed in units of both metres and inter‐plant spaces. In both units, bumblebees progressed more slowly across patchily dispersed arrays. 3. Plant dispersion had a detectable effect on certain components of bees’ individual inter‐plant moves (e.g. move length in metres) but not on others (e.g. mean turning angle). In some cases, the effects of aggregation on the components of individual moves were probably too small to detect statistically and only emerged in their cumulative effect on bees’ progress. 4. Over short sequences of plant visits, bumblebees revisited plants rarely, but the frequency of revisits was almost twice as high in patchy arrays as in uniform arrays, and economic penalties may result from foraging among highly aggregated plants. 5. The effects of plant dispersion on pollinator movements detected in this study are unlikely to have a major impact on pollen transfer in B. napus because differences in pollinator progress only emerged after several successive inter‐plant flights away from a potential pollen source, by which time the limited extent of pollen carryover in this species means that fertilisations from the source plant would be very rare.     

菌根共生网络嵌套性判定的零模型选择

零模型是判定网络嵌套性的重要依据, 菌根共生关系网络经常出现高度非对称性, 该文通过探究矩阵非对称变化对基于不同零模型构建方法的网络嵌套性的影响, 试图为非对称网络零模型的选择提供依据。结果表明: 不同零模型保守性不同, 增加限定条件减少零模型构建过程中的自由空间, 高度限定条件易导致第II类错误。高度非对称网络会增加基于完全随机(r00)零模型的矩阵温度(NT)偏离、降低配对重叠度(NODF)偏离, 标准化指数z-score值显示网络非对称增加后有助于NT和NODF显著性判定。行或列限定对非对称网络嵌套性判定的影响存在差异, 列限定(c0)的网络嵌套性判定对网络非对称性变化的响应规律与r00零模型的响应趋势基本一致, 具有更低的嵌套性偏离和标准差值。行限定(r0, 包括行列限定(backtrack))零模型NT值和NT偏移随矩阵非对称性的变化保持稳定, 较之c0零模型在高度非对称网络中呈现更低的NODF偏离值。选用完全随机和限定零模型相结合的方法, 有助于更加准确判断非对称网络是否具有嵌套结构。高度非对称网络嵌套性判定中对行属性特征比较敏感, 不同非对称性网络间嵌套性水平相比较时选用r0零模型要优于r00和c0零模型。     

明亮熊蜂和意大利蜜蜂在温室桃园的访花行为和传粉生态学比较

2004—2006年连续3年应用明亮熊蜂和意大利蜜蜂在北京为温室桃园传粉,对其访花行为和传粉生态学进行研究.结果表明：两种蜂都可以替代人工掸花为温室桃园提供有效的传粉服务.两种蜂的访花行为和传粉效果不同,明亮熊蜂偏爱于采集花粉,主要以震动翅膀的方式来使花粉释放和传播,而意大利蜜蜂偏爱于采集花蜜,主要以身体接触的方式来粘附和传播花粉;明亮熊蜂的活动起点温度低、日工作时间长、访花速度快,在低温条件下比意大利蜜蜂的传粉效果好;意大利蜜蜂的趋光性强,飞撞温室塑料薄膜的现象严重,受温度和光照条件的影响较大,对温室环境的适应性较差.     

A test of spatial memory and movement patterns of bumblebees at multiple spatial and temporal scales

Naive bumblebee foragers appear to use movement rules at smallspatial and temporal scales, but it is not clear whether theserules determine movement patterns as the scales increase. Onestrategy for efficient foraging used by bumblebees is near-farsearch, involving short flights when in good patches of flowersand longer flights when in poor patches. Bumblebees also demonstratethe use of a spatial memory strategy by returning repeatedlyto patches of flowers, and even following the same route betweenflowers, over periods of days. We attempted to determine atwhat spatial scales bumblebees use spatial memory while foragingwithin a patch and after how many flower visits spatial memoryoutweighs near-far search. Bumblebees in the laboratory foragedon a 4 x 4 array of artificial flowers with distances rangingfrom 10 to 80 cm between flowers in two simple spatial patterns.The proportion of visits to flowers containing a sucrose rewardwas monitored for either 100 or 400 flower visits in two separateexperiments, after which the locations of the rewarding andnonrewarding flowers were interchanged, producing a mirror image.A drop in accuracy after the mirror image switch would indicatethat the bees had memorized the location of rewarding flowers.Mirror image tests, and comparisons to a simulation model ofnear-far search based on actual flight distances, indicate thatnaive bumblebees used near-far search on flowers 10 cm apartbut increasingly used spatial memory as experience and spatialseparation increased. Bumblebees thus have multiple tacticsavailable to forage efficiently in different environments.     

Colony ergonomics for a desert-dwelling bumblebee species (Hymenoptera: Apidae)

Abstract.

• 1 Data on worker traffic, size and concentration of nectar loads, and size and composition of pollen loads were collected for a colony of Bombus pennsylvanicus sonorus Say in the Chihuahuan Desert in Arizona, U.S.A.
• 2 Foraging activity increased through the morning to a peak level in early afternoon and then declined steadily thereafter. Pollen collection occurred primarily in the morning, whereas nectar was harvested throughout the day. Nectar loads decreased in size but increased in sugar concentration during the day.
• 3 Following field observations, we excavated the nest and counted the numbers of immatures and adults present and measured the honey and pollen reserves. A total of 150 workers were present, and we estimate that the colony would have produced 174 queens and 192 males. Food reserves appeared small: pollen and honey stored in the nest represented only 18% and 35%, respectively, of the daily input.
• 4 These results are compared to ergonomic data collected for B.vosnesenskii in the only other similar study.

     

An economic model of the limits to foraging range in central place foragers with numerical solutions for bumblebees

1. A model is described that evaluates the maximum economic foraging range in central place foragers by using optimality criteria to discriminate between foraging sites at different distances from the forager's central place. 2. The basic model can be varied to suit foragers that optimise either their rate of net energy uptake or their foraging efficiency. 3. The model requires specification of the time and energy budgets of travel and foraging, and of the rewards obtainable at potential foraging sites. 4. The specific case of bumblebees, whose foraging ranges are poorly known, is considered. 5. Numerical solutions of the model for parameter values that represent bumblebees and their forage predict economic foraging ranges exceeding several kilometres. The model demonstrates that economics alone can explain extensive flight ranges in bees.     

IS THE PARTIAL MANTEL TEST INADEQUATE?

Abstract.— A publication by Raufaste and Rousset (2001) questioned the effectiveness of the partial Mantel test, a nonparametric statistical test for association among three distance matrices. By repeated simulations, we calculated the cumulative density functions of the null-hypothesis probability of no correlation, within an explicit model of causal relationships. Results do not support the criticism: in conditions of moderate correlation between the independent matrices, the actual error rate is closely associated to the intended type-I error a.     

小峰熊蜂访花偏爱性

传粉昆虫在访花时,通常会表现出对某一类型花的偏爱性。本研究利用人工制作的大小、颜色、形态和气味不同的9种类型的花来研究小峰熊蜂Bombus hypocrita的访花偏爱性。结果表明：当增加花的大小、形态、气味等附加特征数时,小峰熊蜂的偏爱性程度与花朵附加特征数有显著相关性（P<0.01）。当花朵颜色由2种增加到4种,熊蜂对紫色花的偏爱性程度降低,但花朵颜色的种类与小峰熊蜂的访花偏爱性没有相关性（P>0.05）,花颜色的种类对熊蜂访紫色花的偏爱性影响不大。大小为5 cm的紫花被访次数（108±9次）明显高于大小为3 cm的紫花被访次数（40±4次）（P<0.01）,说明熊蜂明显喜欢访大花瓣的紫花。完全盛开的紫花被访次数（129±13次）显著高于刚绽放的花被访次数（26±3次）（P<0.01）,说明熊蜂喜欢访盛开的紫花。柠檬味的紫花被访次数（63±8次）明显低于草莓味的紫色花被访次数（88±2次）（P<0.05）,说明熊蜂喜欢访草莓味的花朵。