Spatial and temporal variation in the provisioning behaviour of female rockhopper penguins Eudyptes chrysocome filholi

As seabirds are central place foragers during breeding, their provisioning behaviour and their ability to face variable energy demand from the chicks is expected to vary with environmental conditions. The provisioning behaviour of female rockhopper penguins Eudyptes chrysocome filholi was recorded over the chick‐rearing period at Kerguelen (KER) and Crozet (CRO) archipelagoes (two very distinct marine environments), using time‐depth recorders, or VHF transmitters coupled with an automatic recording station. No influences of the method have been found on the average foraging trip durations. Some previously undescribed short and multiple trips within a day were recorded using the automatic recording system. These multiple trips (6.8 h) were mostly performed with <5 days old chicks, a period during which feeding rates were the highest (1.1 meals per day), at both sites. During the brooding period, both KER and CRO females mainly performed daily trips of increasing duration (2 h longer at CRO) and at decreasing frequency. During the crèche compared to the brooding period, females from KER performed slightly fewer daily trips (0.6 per day) and more (<3 days) overnight trips, while females from CRO performed very few daily trips (0.1 per day) and more overnight trips, some of them being long trips lasting 5 to 29 days, mostly initiated during the transition between the brooding and the crèche periods. The result fit the hypothesis that long trips permit females to restore and/or maintain their body condition at more distant foraging places. It seemed that chick developement during the brooding period and environmental factors during the crèche period conditioned trip duration of females. Due to more long trips at CRO, the female feeding frequency was twice as high at KER than at CRO during the crèche period, while males participated in the feeding duties. Based on differences in female behaviour, we hypothesize that the male's contribution is likely to differ strongly from one site to another, and may buffer the possible decrease in female feeding frequency by feeding the chicks if food is less abundant.     

Nectar replenishment and pollen receipt interact in their effects on seed production of Penstemon roseus

Resource supply and pollen delivery are often thought to equally limit seed production in animal-pollinated plants. At equilibrium, plants should show no response to experimental pollen supplementation because resources limit seed set above the current level of pollen attraction, while experimental reduction in pollen deposition below the equilibrium level would reduce seed set. The predicted equilibrium may be disrupted, however, if plants expend additional energy to replenish removed nectar. We investigated the combined effects of nectar removal and pollen delivery on female reproductive success of Penstemon roseus (Plantaginaceae), a hummingbird-pollinated plant that replenishes removed nectar. We first documented that the frequency of experimental nectar removal was correlated with total nectar secretion; and increased frequency of nectar removal resulted in increased female reproductive costs to the plant. Trade-offs between investing resources in nectar and investing resources in seeds were then investigated in two contrasting natural populations by removing nectar from flowers at increasing frequencies while simultaneously hand-pollinating flowers with increasing amounts of pollen. Seed set was lowest at low levels of pollen deposition, highest at medium-sized pollen loads, and intermediate when pollen loads were highest. At both sites, the frequency of nectar removal and pollen deposition had an interactive effect on seed production, in that intermediate levels of nectar removal result in the absolute highest seed set, but only at intermediate pollen loads. At high pollen loads, seed set was higher following little to no nectar removal, and at low pollen loads, all rates of nectar removal affected fecundity equally. Seed mass responded to nectar removal and pollination differently than did seed set. High levels of nectar removal and pollen delivery both lowered seed mass, with little interaction between main effects. Our findings are among the first to demonstrate that nectar replenishment costs and pollination intensity jointly affect seed production. This conflict between nectar replenishment costs and pollen-limiting factors results in trade-offs between pollinator attraction and seed provisioning. Thus, resource allocation towards nectar production should more often be considered in future studies of pollen limitation.     

Reproductive biology and effect of nectar robbing on fruit production in Macleania bullata (Ericaceae)

Hummingbird-pollinated flowers are frequently subjected to nectar robbing. In this paper, I examine the impact of nectar robbing on plant reproductive success on a hummingbird-pollinated species. After studying the basic aspects of the floral morphology and reproduction of Macleania bullata (Ericaceae) in a tropical montane wet forest in southwest Colombia, I examined the percent of flowers robbed and the effect of nectar robbery on fruit set. The flowers of this species are typical for plants pollinated by long-bill hummingbirds. They are protandrous and open for four days. Fruit production requires a pollinator visit; fruit set following pollinator exclusion was zero. Fruit set following xenogamous pollen transfer (36.8%) differed significantly from that of population controls (11.9%) and of autogamous pollen transfer (6.3%). Nectar volume, sugar concentration and sugar production were measured at daily intervals from bud opening until the fading of flowers. Daily nectar production (both volume and amount of sugar) varied considerably with flower age. Sugar production peaked on the second day, coinciding with the male phase. The frequency of nectar robbing in the studied population was very high (75% of examined flowers) and was positively correlated with reduced fruit set. I discuss the probability of a relation between reduced fruit set on robbed flowers and an energetic investment. Robbing by non-pollinating visitors can suppose the plant to re-synthesize more nectar. The high incidence of nectar robbing impugns the advantage of specialization.     

Correlation between octopaminergic signalling and foraging task specialisation in honeybees

Regulation of pollen and nectar foraging in honeybees is linked to differences in the sensitivity to the reward. Octopamine (OA) participates in the processing of reward-related information in the bee brain, being a candidate to mediate and modulate the division of labour among pollen and nectar foragers. Here we tested the hypothesis that OA affects the resource preferences of foragers. We first investigated whether oral administration of OA is involved in the transition from nectar to pollen foraging. We quantified the percentage of OA-treated bees that switched from a sucrose solution to a pollen feeder when the sugar concentration was decreased experimentally. We also evaluated if feeding the colonies sucrose solution containing OA increases the rate of bees collecting pollen. Finally, we quantified OA and tyramine (TYR) receptor genes expression of pollen and nectar foragers in different parts of the brain, as a putative mechanism that affects the decision-making process regarding the resource type collected. Adding OA in the food modified the probability that foragers switch from nectar to pollen collection. The proportion of pollen foragers also increased after feeding colonies with OA-containing food. Furthermore, the expression level of the AmoctαR1 was upregulated in foragers arriving at pollen sources compared with those arriving at sugar-water feeders. Using age-matched pollen and nectar foragers that returned to the hive, we detected an upregulated expression of a TYR receptor gene in the suboesophageal ganglia. These findings support our prediction that OA signalling affects the decision in honeybee foragers to collect pollen or nectar.     

Pollination ecology of Penstemon roseus (Plantaginaceae), an endemic perennial shifted toward hummingbird specialization?

Summary Pink-flowered tubular Penstemon roseus (Plantaginaceae), which has shifted partially to hummingbird pollination, blooms on high-elevation slopes in the mountains in Tlaxcala, Mexico. We studied the interactions between pollinator visitation rates to flowers, pollen removal and deposition, flower size, and nectar removal frequency on seed production in P. roseus. We combine observational and experimental studies in two contrasting natural populations. Our manual pollinations revealed that P. roseus is fully self-compatible. Autonomous self- and manual self-pollinated flowers matured as many seeds as when outcrossed, but outcrossing seems to become better than selfing as the flowering season progressed. Early in the season flowers that were bagged and hand-selfed, hand-outcrossed, or autonomously selfed, or unbagged and naturally pollinated had equal seed set in all four treatments. But later in the season, outcross pollen gave approximately twice as much seed set as the two self-treatments. Low levels of pollen receipt and pollen removal were consistent with the long time elapsed for a given plant to be visited by hummingbirds, which suggests pollen shortage in both sites. Despite differences in pollinator visitation rates to flowers, probability of flower visitation, removal and deposition of pollen, and nectar production rates between populations, we found that total nectar production had no effect on seed production at either site. The daily nectar secretion rate of 0.3–0.65 mg sugar per flower per 1–3 days was low relative to other hummingbird-adapted Penstemon species (typical range: 1.5–5 mg sugar per flower), and it might be intermediate between hummingbird- and bee-adapted Penstemon flowers. Our results support the hypothesis about a shift toward hummingbird pollination, and provide an example of a ‘despecialized’ Penstemon species, which attracts high-energy pollinators (hummingbirds) and profits from outcrossing, but retains bee-syndrome floral traits and low sugar production rates.     

Reproductive biology of the melon cactus, Melocactus curvispinus (Cactaceae)

We examined the reproductive biology of Melocactus curvispinus Pfeiffer (Cereeae, Cactoideae) in xeric areas of northern Venezuela. Floral traits correspond to a classic hummingbird-pollination syndrome; however, pollination is shared between hummingbirds, Leucippus fallax (Bourcier 1843), and anthophorid bees, Ceratina sp. Reproduction occurs during most of the year. Anthesis and nectar secretion occur between noon and sunset. Average daily nectar production per flower was 163.1 l, nectar sugar concentration between 29.1 and 30.2% (w/w). Hummingbirds promoted inter-plant pollen movement and were relatively more reliable floral visitors than anthophorid bees, but these insects had a higher frequency of floral visits (28.75/day) than hummingbirds (4.96/day). M. curvispinus is self-compatible and autogamous. By combining extended reproductive activity, frequent animal-mediated pollination, and selfing capabilities, this cactus possesses a mating strategy that guarantees reproduction under variable environmental conditions. We argue that based on its reproductive biology, M. curvispinus should be considered an example of evolutionary transition towards selfing within tribe Cereeae.     

Generalized host‐plant feeding can hide sterol‐specialized foraging behaviors in bee–plant interactions

Host‐plant selection is a key factor driving the ecology and evolution of insects. While the majority of phytophagous insects is highly host specific, generalist behavior is quite widespread among bees and presumably involves physiological adaptations that remain largely unexplored. However, floral visitation patterns suggest that generalist bees do not forage randomly on all available resources. While resource availability and accessibility as well as nectar composition have been widely explored, pollen chemistry could also have an impact on the range of suitable host‐plants. This study focuses on particular pollen nutrients that cannot be synthesized de novo by insects but are key compounds of cell membranes and the precursor for molting process: the sterols. We compared the sterol composition of pollen from the main host‐plants of three generalist bees: Anthophora plumipes, Colletes cunicularius, and Osmia cornuta, as well as one specialist bee Andrena vaga. We also analyzed the sterols of their brood cell provisions, the tissues of larvae and nonemerged females to determine which sterols are used by the different species. Our results show that sterols are not used accordingly to foraging strategy: Both the specialist species A. vaga and the generalist species C. cunicularius might metabolize a rare C₂₇ sterol, while the two generalist species A. plumipes and O. cornuta might rather use a very common C₂₈ sterol. Our results suggest that shared sterolic compounds among plant species could facilitate the exploitation of multiple host‐plants by A. plumipes and O. cornuta whereas the generalist C. cunicularius might be more constrained due to its physiological requirements of a more uncommon dietary sterol. Our findings suggest that a bee displaying a generalist foraging behavior may sometimes hide a sterol‐specialized species. This evidence challenges the hypothesis that all generalist free‐living bee species are all able to develop on a wide range of different pollen types.     

Population dynamics of the solitary digger bee<Emphasis Type="Italic"> Andrena vaga</Emphasis> Panzer (Hymenoptera,Andrenidae) studied using mark-recapture and nest counts

The population size of the solitary digger bee Andrena vaga was documented using mark-recapture methods during 4 consecutive years (1996–1999). Additionally, the numbers of the parasitoids Nomada lathburiana and Bombylius major were estimated. For calculations of the daily population size, the Jolly-Seber model for open populations with time-dependent survival and capture rates was used. The mean daily population size corresponded well to the number of nests marked in the aggregation of A. vaga. The method of mark-recapture achieved the most accurate data when the number of sampling periods and the capture probability was high. The results document a decrease in population size of the investigated A. vaga aggregation during 4 successive years. A population increase of its parasitoids B. major is considered to be one reason for the population decline of A. vaga in 1999. The sex ratio of A. vaga was biased toward females, contrary to the expected higher number of males. Results of the mark-recapture studies showed that up to 50% of females left the aggregation or died at the beginning of the season. The females probably colonized new aggregations at distances further than 200 m away from their birth aggregation, as no marked females were observed in the environs of the study area. This behavior is interpreted as a parasite avoidance strategy. The bee populations in the study area form a web of interconnected aggregations. Apparently a flux of extinction and recolonization exists between the different aggregations.     

Floral nectar sugar composition and flowering phenology of the food plants used by the western pygmy possum, <Emphasis Type="Italic">Cercartetus concinnus</Emphasis>, at Innes National Park,South Australia

The western pygmy possum (Cercartetus concinnus) is a small nocturnal marsupial that relies primarily on the nectar and pollen of myrtaceous species at Innes National Park and may occasionally also ingest invertebrates. This study confirmed plant utilization by C. concinnus using scat samples and pollen swabs, and investigated the flowering phenology of dietary plants to determine resource availability. We compared nectar composition between day and night and analyzed nectar sugar production for dietary species. Pollen swabs and scats suggested that C. concinnus relied primarily on the nectar and pollen of Kingscote mallee, Eucalyptus rugosa (76.8% of grains counted in combined scat samples) at Innes National Park, when available; only one of 30 scat samples contained numerous moth scales. The nectars of the species investigated showed marked differences in their composition, but only Melaleuca gibbosa and M. halmaturorum sugar composition changed between day and night. The nectar sugar ratio of E. rugosa differed from those of most other species investigated. C. concinnus may select this plant’s flowers because its nectar is relatively high in hexose sugars. Although E. diversifolia was abundant, its flowers were mostly ignored by possums, perhaps because the nectar in these flowers was proportionately much richer in sucrose than other species’. E. rugosa’s flowering index (calculated from flower load and canopy size) was greatest in December. Six of the seven eucalypt species flowered between November and April; for half of the year pygmy possums must find other resources.     

Windows of opportunity and the temporal structuring of foraging activity in a desert solitary bee

1. Females of the desert solitary bee Anthophora pauperata collect nectar and pollen almost exclusively from Alkanna orientalis (Boraginaceae). The bee and plant are found together in the early spring, living in the bottom of steep-sided wadis (dry river valleys) at an altitude of 1500 m in Egyptian Sinai. 2. Female A. pauperata showed clear morning and afternoon peaks in foraging activity, separated by a 2–3 h midday period spent in their underground nests. This study analyses the following in order to identify the factors structuring this daily pattern: thermal aspects of the bee and its environment, temporal patterns of resource provision by the plant, and female nectar and pollen foraging behaviour. 3. Although A. pauperata can generate substantial heat endothermically, morning and evening ambient temperatures well below 10 °C defined a thermal window within which foraging occurred. Maximum air temperatures were moderate (25–30 °C), and examination of the physiology and behaviour of A. pauperata suggests that the midday reduction in flight activity was not due to thermal constraints. 4. Alkanna orientalis produces protandrous hermaphroditic flowers. Female A. pauperata collected pollen from male-phase flowers and harvested nectar preferentially from female-phase flowers. Although the nectar standing crop was relatively constant throughout the day, pollen availability peaked strongly in the early afternoon. 5. Female A. pauperata visited young male-phase flowers as soon as they opened, generating an early afternoon peak in pollen foraging activity and depleting the pollen standing crop rapidly. A morning peak in pollen foraging occurred when females gleaned remnant pollen from flowers that had opened the previous day. Pollen availability in the morning was far lower than in the early afternoon, and the time taken to collect a full pollen load in the morning was significantly longer. Collection of pollen in the morning despite very low resource availability suggests that pollen may be a limiting resource for A. pauperata. 6. In contrast to many existing examples of bimodal activity patterns in highly endothermic bees, the bimodal activity patterns of female A. pauperata appear to be driven not by thermal considerations but by daily patterns of pollen release from its principal food source.     

Nectar secretion pattern of the dish-shaped flower,Cayratia japonica (Vitaceae), and nectar utilization patterns by insect visitors

We studied the relationship between the diurnal nectar secretion pattern of flowers of Cayratia japonica and insect visiting patterns to these flowers. Flower morphology of C. japonica changed greatly for about 12 hours after flower-opening and the maximum duration of nectar secretion was 2 days. The nectar volume peaked at 11∶00 and 15∶00, and declined at night and at 13∶00 regardless of time elapsed after flower-opening. The nectar volume at the two peaks was, on average, 0.25 μl on bagged inflorescences and 0.1μl on unbagged inflorescences (both, sugar concentration=60%). The flower secreted nectar compensatory when the nectar was removed. This means that insects consume more nectar than the difference of nectar volume between bagged and unbagged flowers. Apis cerana is a primary visitor of this flower, and was the only species for which we confirmed pollen on the body, among many species of flower visiting insects to this flower. Apis cerana visited intensively at the two peaks of nectar secretion. Visits of the other insects were rather constant or intensive only when there was no nectar secretion. Thus flowers of C. japonica with morphologically unprotected nectaries may increase likelihood that their nectar is used by certain pollinators, by controlling the nectar secretion time in day. In this study the pattern of nectar secretion allowed A. cerana maximum harvest of nectar.     

The influence of nectar production on hawkmoth behavior,self pollination,and seed production in Mirabilis multiflora (Nyctaginaceae)

Nectar is the most common floral reward that plants produce to attract pollinators. To determine the effect of nectar production on hawkmoth behavior, pollen movement, and reproductive success in Mirabilis multiflora, I manipulated nectar volumes and observed the subsequent foraging behavior of the hawkmoth Hyles lineata and the resulting pollen movement patterns. Individual hawkmoths visited significantly more flowers on plants with more nectar. The increase in flower visits significantly increased pollen deposition on stigmas and pollen removal from anthers when nectar volume was raised to twice the highest level found in nature. As hawkmoths visited flowers consecutively on a plant, the proportion of self pollen deposited on stigmas increased significantly and rapidly. Based on simulated hawkmoth visits, seed set was significantly reduced for flowers later in a visit sequence. A simple model combining these results predicts that the form of selection on nectar production varies depending on pollinator abundance. Using a multiple regression analysis a nearly significant (P < 0.08) effect of stabilizing selection was detected during a single season as predicted by the model for the prevailing hawkmoth abundance. Although increased nectar production may indirectly affect plant fitness by reducing resources available for other plant functions, the direct effect of high nectar production on pollinator behavior and self pollination may generally limit floral nectar production.     

Tactile learning and the individual evaluation of the reward in honey bees (Apis mellifera L.)

Using the proboscis extension response we conditioned pollen and nectar foragers of the honey bee (Apis mellifera L.) to tactile patterns under laboratory conditions. Pollen foragers demonstrated better acquisition, extinction, and reversal learning than nectar foragers. We tested whether the known differences in response thresholds to sucrose between pollen and nectar foragers could explain the observed differences in learning and found that nectar foragers with low response thresholds performed better during acquisition and extinction than ones with higher thresholds. Conditioning pollen and nectar foragers with similar response thresholds did not yield differences in their learning performance. These results suggest that differences in the learning performance of pollen and nectar foragers are a consequence of differences in their perception of sucrose. Furthermore, we analysed the effect which the perception of sucrose reward has on associative learning. Nectar foragers with uniform low response thresholds were conditioned using varying concentrations of sucrose. We found significant positive correlations between the concentrations of the sucrose rewards and the performance during acquisition and extinction. The results are summarised in a model which describes the relationships between learning performance, response threshold to sucrose, concentration of sucrose and the number of rewards. Accepted: 14 April 1999     

Extrafloral‐nectaries and interspecific aggressiveness regulate day/night turnover of ant species foraging for nectar on Bionia coriacea

Plants bearing extrafloral nectaries (EFNs) vary the secretion of nectar between day and night, which creates turnover in the composition of interacting ant species. Daily variation in the composition of ant species foraging on vegetation is commonly observed, but its mechanisms are poorly understood. We evaluated the daily variation in nectar availability and interspecific aggressiveness between ants as possible regulatory mechanisms of the turnover in ant–plant interactions. We hypothesized that (i) plants would interact with more ant species during periods of higher secretion of nectar and that (ii) aggressive ant species would compete for nectar, creating a daily turnover of species collecting nectar. We tested this hypothesis by measuring the production of nectar during the day and night and by experimentally removing EFNs of Bionia coriacea (=Camptosema coriaceum) (Nees & Mart.) Benth. (Fabaceae: Faboideae) plants in a Brazilian savanna (Cerrado). We then compared the abundance and composition of ant species between those treatments and during the day. Our results indicate that more ant workers forage on plants during the day, when nectar was sugary, while more ant species forage at night, when aggressiveness between ant species was lower. We also detected a day/night turnover in ant species composition. Ant species foraging for nectar during the day were not the same at night, and this turnover did not occur on plants without EFNs. Both dominant ant species, diurnal Camponotus crassus (Hymenoptera: Formicidae) and nocturnal Camponotus rufipes (Hymenoptera: Formicidae), were the most aggressive species, attacking other ants in their specific periods of forage while also being very aggressive toward each other. However, this aggressiveness did not occur in the absence of nectar, which allowed non‐aggressive nocturnal ant species to forage only during the daytime, disrupting the turnover. We conclude that extrafloral‐nectar presence and interspecific aggressiveness between ants, along with other environmental factors, are important mechanisms creating turnovers in ants foraging on plants.     

Field energetics and the estimation of pollen and nectar intake in the marsupial honey possum, Tarsipes rostratus, in heathland habitats of South-Western Australia

A method is described, based on the simultaneous turnover of both stable (¹⁸O) and radioactive isotopes (³H and ²²Na), whereby the daily nectar and pollen intake of free-ranging marsupial honey possums (Tarsipes rostratus) may be estimated. The field metabolic rate is measured using doubly labelled water and nectar intake is estimated independently from the measured water and sodium fluxes. The method assumes that free-water intake is negligible (but may be accounted for if not the case), that virtually all dietary sodium is derived from nectar rather than from pollen, and that the animals are in energetic balance over the period of measurement. These assumptions have been tested and found to be robust, except during periods of heavy rain when significant intakes of free-water were recorded. Leaching experiments with pollen grains suggest that less than 10% of the sodium ingested by honey possums is derived from pollen and calculations thus assumed a 90%:10% split between nectar and pollen. Nectar intake averaged 5.9 ± 0.6 ml · day⁻¹ and regressing nectar intake on daily change in body mass predicts an intake of approximately 7 ml · day⁻¹ nectar to maintain balance for a 9 g honey possum. Estimates of pollen intake averaged 660 ± 156 mg · day⁻¹ and a similar regression analysis of the data predicts that a daily intake of approximately 1 g pollen would be needed to maintain mass balance of honey possums. Estimated nectar and pollen intakes did not differ significantly between males and females, but nectar intake was higher in winter compared with dry periods of the year. The sugar content of nectar falls during winter, however, and the overall energy derived from nectar thus remains roughly constant. Estimates of pollen and nectar intake for individual animals were not significantly correlated, suggesting that honey possums forage selectively for these two food items. Accepted: 19 August 1999     

Foraging behavior of honeybees on white clover

Summary Measurements of several aspects of foraging behavior of honeybees on white clover and of factors that might influence it were made at one-to-two-week intervals during the season. Other measurements were made on intervening days. Individual blossoms yielded a mean of from 0.02 to 0.08 l of nectar containing 42 to 65% sugar during the season. On one day when the nectar averaged 65% sugar, bees with pollen on their corbiculae carried a mean of 5.2 mg of pollen and 37.3 mg of nectar; bees without pollen carried a mean of 37.9 mg of nectar. On 10 occasions each of 50 bees were timed while foraging a single blossom; the mean foraging speed differed significantly on the 10 occasions, and varied from 1.5 to 3.0 seconds per blossom. Each of 18 bees was timed during visits to 25 blossoms, and the total time required for the same bees to forage from the blossoms on 25 racemes was also measured. The mean foraging speed of these bees varied from 1.1 to 4.5 seconds per blossom; the differences were highly significant. These bees foraged 7.0 to 26.0 blossoms per minute, and there was a strong negative correlation between the mean seconds required for bees to forage blossoms and the mean number of blossoms foraged per minute. Fifty one bees observed for a total of 314.5 minutes foraged an average of 18.4 blossoms per minute and 3.4 blossoms per inflorescence. Sixty one per cent of the flights of foraging bees were made to inflorescences estimated to be less than 5 inches from the starting point.Published as Technical Contribution N^o 4244 from theTexas Agricultural Experiment Station.     

Food provisioning and stone handling tradition in Japanese macaques: a comparative study of ten troops

By addressing the influence of food provisioning on stone handling (SH), a behavioral tradition in Japanese macaques, this study contributes to the ongoing debate in cultural primatology by asking whether human intervention influences the emergence or propagation of behavioral traditions. SH is a form of object play consisting of the manipulation of stones by performing various behavioral patterns. We tested the hypothesis that the frequency of food provisioning affects the daily performance, form, and context of occurrence of SH by influencing a troop's feeding-related activity budget. We used a standardized observation procedure to investigate SH in ten troops of Japanese macaques. In troops provisioned several times a day, SH was more frequent, longer, and more prevalent during provisioning than nonprovisioning periods. These effects of provisioning were not significant in troops provisioned less frequently. SH was more frequently integrated with food-related activities in troops supplied with food several times a day than in the other troops. Food provisioning may be a key factor in the innovation and transformation phases of the SH tradition in Japanese macaques.     

Spatial foraging patterns of the African honey bee,Apis mellifera scutellata

Recruitment patterns were investigated for the African honey bee in the Okavango River Delta, Botswana. The waggle dances of two observation colonies maintained in the field were monitored and used to construct maps of daily recruitment activity. These maps revealed that the African colonies frequently adjusted the allocation of recruits among food patches, recruited for 16–17 different food sites/day over areas of 55–80 km ² ,and concentrated the majority of recruitment within 1 km of the hives (median foraging distances for the two colonies were 295 and 563 m). In both colonies pollen foragers were more abundant than nectar foragers, and pollen sources indicated by waggle dancers were significantly closer to the hives than nectar sources. Compared to the recruitment patterns of temperate climate colonies, the African colonies had smaller recruitment areas, smaller mean recruitment distances, and a greater emphasis on pollen foraging. These differences may be related to the contrasting survival strategies followed by tropical-versus temperate-climate honey bees.     

A trade-off between the amount and distance of pollen dispersal triggered by the mixed foraging behaviour of Sephanoides sephaniodes (Trochilidae) on Lapageria rosea (Philesiaceae)

Nectar thieves may increase or decrease pollinator-mediated pollen flow and thus may have positive or negative effects on plant reproductive success. In temperate rainforests of South America, the hummingbird Sephanoides sephaniodes acts as both a pollinator and non-destructive nectar thief on Lapageria rosea. Although pollinators that also act as nectar thieves have the potential to significantly modify plant reproductive success, no previous study has addressed this. To determine how the mixed behaviour of S. sephanoides affects pollen flow, we experimentally exposed some flowers to nectar theft and excluded nectar thieves from other flowers. We then assessed pollen dispersal into the floral neighbourhood. Thieved flowers exported less pollen, but the pollen exported was transferred farther into the neighbourhood. Our findings indicate a trade-off between distance and amount of pollen flow.     

Pollinator genetics and pollination: do honey bee colonies selected for pollen‐hoarding field better pollinators of cranberry Vaccinium macrocarpon?

1. Genetic polymorphisms of flowering plants can influence pollinator foraging but it is not known whether heritable foraging polymorphisms of pollinators influence their pollination efficacies. Honey bees Apis mellifera L. visit cranberry flowers for nectar but rarely for pollen when alternative preferred flowers grow nearby. 2. Cranberry flowers visited once by pollen‐foraging honey bees received four‐fold more stigmatic pollen than flowers visited by mere nectar‐foragers (excluding nectar thieves). Manual greenhouse pollinations with fixed numbers of pollen tetrads (0, 2, 4, 8, 16, 32) achieved maximal fruit set with just eight pollen tetrads. Pollen‐foraging honey bees yielded a calculated 63% more berries than equal numbers of non‐thieving nectar‐foragers, even though both classes of forager made stigmatic contact. 3. Colonies headed by queens of a pollen‐hoarding genotype fielded significantly more pollen‐foraging trips than standard commercial genotypes, as did hives fitted with permanently engaged pollen traps or colonies containing more larvae. Pollen‐hoarding colonies together brought back twice as many cranberry pollen loads as control colonies, which was marginally significant despite marked daily variation in the proportion of collected pollen that was cranberry. 4. Caloric supplementation of matched, paired colonies failed to enhance pollen foraging despite the meagre nectar yields of individual cranberry flowers. 5. Heritable behavioural polymorphisms of the honey bee, such as pollen‐hoarding, can enhance fruit and seed set by a floral host (e.g. cranberry), but only if more preferred pollen hosts are absent or rare. Otherwise, honey bees' broad polylecty, flight range, and daily idiosyncrasies in floral fidelity will obscure specific pollen‐foraging differences at a given floral host, even among paired colonies in a seemingly uniform agricultural setting.