Foraging behavior of individual workers and foraging dynamics of colonies of three Sumatran stingless bees

The foraging behavior of three stingless bees, Trigona (Tetragonula) minangkabau, T. (Trigonella) moorei and T. (Heterotrigona) itama, was studied to describe patterns of resource harvest in disturbed forest areas in Sumatra, Indonesia.

1. Average daily total number of foraging flights per colony was 1200 in T. minangkabau, 2400 in T. moorei and 7000 in T. itama and it was proportional to colony population size. Foragers collecting nectar, pollen or plant resin were respectively 70–80%, 10–20% and <10% in the three species. Pollen was collected most in the morning. Nectar collection peaked in midday in T. itama but continued almost evenly until dusk in T. minangkabau and T. moorei. Resin was collected evenly throughout day.
2. In all the three species the volume of a nectar load carried by a returning forager did not decrease until 1600, followed by slight decline. In the morning the sugar concentration of nectar was almost constantly 30%, and in the afternoon its maximum value reached 60% although some remained 30%. As a result, the mean sugar weight in a nectar load gradually increased until dusk.
3. In T. minangkabau, resin was collected by specialized foragers. Nectar foragers switched to collect pollen and vice versa. There were two types of foraging: “exploitatory” flights, the repitition of short, rather constant flights bringing back full resource loads, and “exploratory” flights, prolonged flight and/or reduction in amount of resources carried. Exploitatory flights followed the exploratory flights which led to discovery of rich sources. Mean duration of exploitatory flights was 7 min during nectar collection, 12 min for pollen and 23 min for resin. Sites of exploited flowers were estimated to be 84–434 m distant from the nest site. Mean duration of flights for nectar collection was 13 min in the morning and 6 min in the late afternoon.

     

Activity patterns of females of the solitary bee Anthophora plumipes in relation to temperature, nectar supplies and body size

Abstract.

• 1 Patterns of activity at a large nesting aggregation and at foraging sites are described for females of the solitary bee Anthophora plumipes (Pallas). Changes in the quality and quantity of the resource collected by females provisioning cells are related to variation in female body mass and microclimate.
• 2 Activity at the nest site demonstrated relationships with aspects of the thermal environment experienced by A.plumipes. Measures of temperature showing significant relationships differed for females in different stages in the nesting cycle exhibiting characteristically different behaviour patterns.
• 3 Larger females emerged from nest tunnels earlier in the morning and provisioned cells at lower ambient temperatures than smaller individuals. Body size therefore predicts reproductive success at low ambient temperatures.
• 4 Pollen and nectar loads carried by females increased with ambient temperature. Because only one cell is completed per day and the size of offspring is determined by the quality and quantity of resource provided by the mother, the body size of individuals emerging in the following season will depend on interactions between climate and body size, in addition to any heritable component.
• 5 Variation in activity levels at foraging sites is attributed not only to thermal considerations, but also to variation in the quality of rewards available at different floral sources.

     

The relationship between population size,amount of brood,and individual foraging behaviour in the honey bee,Apis mellifera L.

This study experimentally examines the relationship between colony state and the behaviour of individual pollen and nectar foragers in the honey bee, Apis mellifera L. In the first experiment we test the prediction that individual pollen foragers from colonies with higher brood quantities should exhibit a greater work effort for pollen resources than individual pollen foragers from colonies with low brood quantities. Eight colonies were assigned into two treatment groups; HIGH brood colonies were manipulated to contain 9600±480 cm² brood area; LOW brood colonies were manipulated to contain 1600±80 cm² brood area. We measured colony brood levels over the course of the experiment and collected individual pollen loads from returning pollen foragers. We found that, while colonies remained significantly different in brood levels, individual pollen foragers from HIGH brood colonies collected larger loads than individuals from LOW brood colonies. In the second experiment we investigated the influence of colony size on the behaviour of individual nectar foragers. We assigned eight colonies to two treatment groups; LARGE colonies were manipulated to contain 35000±1700 adult workers with 3500±175 cm² brood area, and SMALL colonies were manipulated to contain 10000±500 adult workers with 1000±50 cm² brood area. We observed foraging trips of individually marked workers and found that individuals from LARGE colonies made longer foraging trips than those from SMALL colonies (LARGE: 1666.7±126.4 seconds, SMALL: 1210.8±157.6 seconds), and collected larter nectar loads (LARGE: 19.2±1.0 l, SMALL: 14.6±0.8 l). These results indicate that individual nectar foragers from LARGE colonies tend to work harder than individuals from SMALL colonies. Both experiments indicate that the values of nectar and pollen resources to a colony change depend on colony state, and that individual foragers modify their behaviour accordingly.     

Pollen of Apis honey from Maharashtra

Melissopalynological investigations are important in establishing an apiary industry. Pollen analyses of the nectar sources of Apis florea Fabr. from the Saoner region indicate the importance of this region for commercial bee keeping enterprises. Twenty seven samples were collected during winter season in the months of October 1997 to January 1998 and November 1998 to January 1999. In addition, 245 pollen loads were analyzed from 19 honey combs collected from the same region. Pollen grains were identified from reference slides. Of the 27 honey samples, 17 samples were unifloral and ten multifloral. A total of 39 pollen types were classified into 21 families. In the analyses of the 245 pollen loads Lagascea mollis Cav. was dominant in 57 pollen loads and Parthenium hysterophorus L. in 55 pollen loads. Weeds such as Parthenium hysterophorus, Sphaeranthus indicus L., Lagascea mollis, Bidens pilosa L. and Justicia procumbens L. were the common sources of nectar and pollen for Apis florea in the region. Agricultural tracts of the Saoner region in the Nagpur District have a good potential for sustaining beekeeping ventures because of the diversity of nectar and pollen taxa.     

Specialised protagonists in a plant–pollinator interaction: the pollination of Blumenbachia insignis (Loasaceae)

• Analyses of resource presentation, floral morphology and pollinator behaviour are essential for understanding specialised plant‐pollinator systems. We investigated whether foraging by individual bee pollinators fits the floral morphology and functioning of Blumenbachia insignis, whose flowers are characterised by a nectar scale‐staminode complex and pollen release by thigmonastic stamen movements.
• We described pollen and nectar presentation, analysed the breeding system and the foraging strategy of bee pollinators. We determined the nectar production pattern and documented variations in the longevity of floral phases and stigmatic pollen loads of pollinator‐visited and unvisited flowers.
• Bicolletes indigoticus (Colletidae) was the sole pollinator with females revisiting flowers in staminate and pistillate phases at short intervals, guaranteeing cross‐pollen flow. Nectar stored in the nectar scale‐staminode complex had a high sugar concentration and was produced continuously in minute amounts (~0.09 μl·h^?1). Pushing the scales outward, bees took up nectar, triggering stamen movements and accelerating pollen presentation. Experimental simulation of this nectar uptake increased the number of moved stamens per hour by a factor of four. Flowers visited by pollinators received six‐fold more pollen on the stigma than unvisited flowers, had shortened staminate and pistillate phases and increased fruit and seed set.
• Flower handling and foraging by Bicolletes indigoticus were consonant with the complex flower morphology and functioning of Blumenbachia insignis. Continuous nectar production in minute quantities but at high sugar concentration influences the pollen foraging of the bees. Partitioning of resources lead to absolute flower fidelity and stereotyped foraging behaviour by the sole effective oligolectic bee pollinator.

     

Plant Resources Foraged by Polybia Scutellaris (Hym. Vespidae) in the Argentine Pampas

The plants foraged by Polybia scutellaris were studied by analysing the pollen of honey collected from six nests. 34 morphological types were identified. These honeys showed a predominance of Poaceae and Cyperaceae pollen. The main access to anemophilous pollen is through the honeydew produced by Claviceps in Paspalum dilatatum. The principal sources of nectar were the native “Astereae” (Aster squamatus, Baccharis sp. and Solidago chilensis) and Eucalyptus sp. When collecting honeydew and nectar, wasps foraged plants close to their nest. Most of the plants visited belonged to a dense population. The successive flowering reflected in these honeys, shows that wasps also consume honey in summer and that it is not all stored for the winter season.     

Colony pollen reserves affect body size,sperm production and sexual development in males of the stingless bee <Emphasis Type="Italic">Melipona beecheii</Emphasis>

The production of male sexual offspring by social insect colonies is often strongly seasonal or resource-dependent. In stingless bees, males are produced in smaller numbers under conditions of low colony food reserves; whether such males are negatively affected in traits related to reproductive success is not known. We compared body size, sperm production and sexual maturity in Melipona beecheii males reared with experimentally supplemented or reduced pollen reserves, but with otherwise equal numbers of workers and equal quantities of honey reserves. We also studied the same traits in males collected from non-manipulated colonies with pollen reserves intermediate between the supplemented or reduced groups but with more workers and honey reserves. Males reared under experimentally reduced pollen reserves had significantly smaller bodies and lower sperm counts compared to those reared in colonies with experimentally supplemented pollen reserves. There was also a significantly positive relationship between the number of sperm and body size in males across all colony treatments. The maximum number of sperm in seminal vesicles was recorded 2 days later in males from colonies with reduced pollen compared to males from colonies with supplementary pollen. Males from non-manipulated colonies were intermediate in size, sperm count and speed of maturation. Our study documents for the first time the existence of large size variation in males of stingless bees that is related with the amount of pollen reserves in their natal colony. We conclude that a colony’s pollen reserves have a major impact on male body size, sperm production and speed of sexual maturity in this stingless bee, which may be the case in other social insects. Stingless bees are a good model system to study the balance between colony-level selection and individual-level selection on male sexually selected traits such as body size.     

The superiority of bumblebees to honeybees as pollinators: insect visits to raspberry flowers

Abstract.

• 1 The behaviour and activity patterns of Apis mellifera and of five species of Bombus were analysed in relation to climatic variables and nectar quality on three varieties of unsprayed cultivated raspberry (Rubus idaeus) in eastern Scotland.
• 2 Stages of floral morphology and reward were similar for the three varieties: young flowers offered both nectar and pollen, but medium and old flowers offered nectar only, in diminishing quantities.
• 3 A wide range of insects visited raspberry flowers, but bees were dominant, bumblebees being responsible for about 60% of all visits and honeybees making up most of the remaining percentage. All bees had substantial pollen deposited on their bodies during visits, though few specifically collected it.
• 4 Bombus spp. were found to favour young (receptive) flowers strongly, especially early in the morning when pollen was most abundant: whilst Apis visited unselectively. Bumblebees also foraged over substantially longer periods of the day, and in poorer weather, some being present at most times of observation; and they foraged more quickly in terms of flower visits per minute.
• 5 Bombus carried more pollen on their bodies than Apis, and also deposited more pollen on raspberry stigmas, with B.lapidarius and B.terrestris being particularly effective and also being the most abundant species. All bumblebees also foraged over a longer range, moving between canes and rows more frequently than did honeybees.
• 6 Bumblebees are therefore likely to be substantially more important as pollinators of raspberries than are honeybees, especially as raspberries though moderately self-fertile may exhibit metaxenia. Reasons why Bombus may be the preferred pollinator in most sites of raspberry cultivation are discussed, together with implications for present and future growers.

     

Evidence for mutualism between a flower-piercing carpenter bee and ocotillo: use of pollen and nectar by nesting bees

Abstract.

• 1 Carpenter bees (Xylocopa californica arizonensis) in west Texas, U.S.A., gather pollen and ‘rob’ nectar from flowers of ocotillo (Fouquieria splendens). When common, carpenter bees are an effective pollen vector for ocotillo. We examined ocotillo's importance as a food source for carpenter bees.
• 2 The visitation rate of carpenter bees to ocotillo flowers in 1988 averaged 0.51 visits/flower/h and was 4 times greater than that of queen bumble bees (Bombus pennsylvanicus sonorus), the next most common visitor. Nectar was harvested thoroughly and pollen was removed from the majority of flowers. Hummingbird visits were rare.
• 3 Pollen grains from larval food provisions were identified from sixteen carpenter bee nests. On average, 53% of pollen grains sampled were ocotillo, 39% were mesquite (Prosopis glandulosa), and 8% were Zygophyllaceae (Larrea tridentata or Guaiacum angustifolium). Carpenter bee brood size averaged 5.8 per nest.
• 4 We measured the number of flowers, and production of pollen and nectar per flower by mature ocotillo plants, as well as the quantity of pollen and sugar in larval provisions. An average plant produced enough pollen and nectar sugar to support the growth of eight to thirteen bee larvae. Ocotillo thus has the potential to contribute significantly to population growth of one of its key pollinators.
• 5 Although this carpenter bee species, like others, is a nectar parasite of many plant species, it appears to be engaged in a strong mutualism with a plant that serves as both a pollen and as a nectar source during carpenter bee breeding periods.

     

Floral resources foraged by Geotrigona argentina (Apidae,Meliponini) in the Argentine Dry Chaco forest

This study is the first contribution to knowledge of the relationships between Geotrigona argentina and the plants of the Argentine Dry Chaco forest. A total of 1260 g of honey (corresponding to 146 pots) and 763 g of pollen (63 pots) stored in four underground nests was studied. The honey pots from each nest were homogenised and the four honey samples were analysed by melissopalynological methods, whereas the pollen pots were studied individually. Both classical counts and counts affected by the volume of the pollen types were carried out. Pollen data were statistically analysed. Additional data on both protein and lipid content is also provided. A total of 39 pollen taxa were identified. Pollen collection was focused on a few pollen taxa: Prosopis, Castela coccinea, Maytenus and Capparis; these taxa, together with Ziziphus mistol and Pisonia zapallo, were also important nectar sources. The preliminary results show that pollen collection varied seasonally, being most diverse in the summer when G. argentina incorporates herbaceous plants into its diet. The pollen collection spectrum of G. argentina is similar to that of other Trigonina bees in that the main plant species collected are a few large shrubs or trees, whose flowering consists of small and clustered flowers. Pots with large amounts of monofloral loads with pollen from only a few species suggests an organised foraging behaviour that includes the recruitment of foragers, such as that observed in other eusocial bees.     

Floral preferences and climate influence in nectar and pollen foraging by Melipona rufiventris Lepeletier (Hymenoptera: Meliponini) in Ubatuba, São Paulo state, Brazil

We describe the environment effects on the amount and quality of resources collected by Melipona rufiventris Lepeletier in the Atlantic Forest at Ubatuba city, S?o Paulo state, Brazil (44o48'W, 23o22'S). Bees carrying pollen and/or nectar were captured at nest entrances during 5 min every hour, from sunrise to sunset, once a month. Pollen loads were counted and saved for acetolysis. Nectar was collected, the volume was determined and the total dissolved solids were determined by refractometer. Air temperature, relative humidity and light intensity were also registered. The number of pollen loads reached its maximum value between 70% and 90% of relative humidity and 18oC and 23oC; for nectar loads this range was broader, 50-90% and 20-30oC. The number of pollen loads increased as relative humidity rose (rs = 0.401; P < 0.01) and high temperatures had a strong negative influence on the number of pollen loads collected (rs = -0.228; P < 0.01). The number of nectar loads positively correlated with temperature (rs = 0.244; P < 0.01) and light intensity (rs = 0.414; P < 0.01). The percentage of total dissolved solids (TDS) on nectar loads positively correlated with temperature and light intensity (rs = 0.361; P < 0.01 and rs = 0.245; P < 0.01), negatively correlated with relative humidity (rs = -0.629; P < 0.01), and it increased along the day. Most nectar loads had TDS between 11% and 30%, with an average of 24.7%. The volume measures did not show any pattern. Important pollen sources were Sapindaceae, Anacardiaceae, Rubiaceae, Arecaceae, Solanaceae and Myrtaceae; nectar sources were Sapindaceae, Fabaceae, Rubiaceae, Arecaceae and Solanaceae.     

Regulation of nectar collection in relation to honey storage levels by honey bees, Apis mellifera

Honey bees collect distinct nutrient sources in the form ofnectar (energy) and pollen (nitrogen). We investigated the effectof varying energy stores on nectar and pollen foraging. We foundno significant changes in nectar foraging in response to changesin honey storage levels within colonies. Individual foragersdid not vary activity rates or nectar load sizes in responseto changes in honey stores, and colonies did not increase nectarintake rates when honey stores within the hive were decreased.This result contrasts with pollen foraging behavior, which isextremely sensitive to colony state. Our data show that individualforaging decisions during nectar collection and colony regulationof nectar intake are distincdy different from pollen foraging.The behavior of honey bees illustrates that foraging strategy(and therefore foraging models) can incorporate multiple currencies,including both energy and protein intake.[Behav Ecol 7: 286–291(1996)]     

Nectar utilization and pollination by Australian honeyeaters and insects visiting Calothamnus quadrifidus (Myrtaceae)

Nectar availability in Calothamnus quadrifidus flowers was studied at Wongamine Nature Reserve in late spring (November). Despite some overnight depletion by moths and other invertebrates, more nectar was present in flowers at dawn than at the preceding dusk. Significant nectar depletion occurred within a few hours after dawn, mainly due to foraging by two honeyeater species. Lichmera indistincta and Phylidonyris nigra. Thereafter, nectar availability was maintained at relatively low levels, principally because of foraging by honeyeaters and honey bees. Apis mellifera, that became active during the warmer part of the day. Although individual honeyeaters consumed more nectar than A. mellifera, honey bees were so abundant that their total impact was greater than that of either honeyeater species for much of the day. Transfer of C. quadrifidus pollen between flowers is necessary in order to achieve a high level of seed set, as the flowers appear to be protandrous. Honeyeaters appeared to be considerably more significant pollen vectors than A. mellifera.     

The giant willow aphid (Tuberolachnus salignus) and its effects on the survival and growth of willows

1. The giant willow aphid (Tuberolachnus salignus) is an invasive pest that can attain large populations on willows (Salix spp.). This has the potential to have a negative impact on the extensive use of willows for soil conservation, and as a source of pollen and nectar for honey bees in New Zealand.
2. A willow nursery field trial was established to evaluate the aphid populations, and the survival and growth of young plants of several willow species and hybrids, during two growing seasons from planting.
3. The willow species and hybrids varied widely in their susceptibility to the aphid, with large aphid populations and plant mortality in the most susceptible willows, and reductions in plant growth in all but aphid-resistant willows. The effects on the plants were not seen in the first season, but occurred during the second season.
4. The aphid can be expected to have some negative impacts in New Zealand, with reductions in growth of some willows commonly used for soil conservation, and for pollen and nectar for honey bees.

     

Foraging behavior of Melipona rufiventris Lepeletier (Apinae; Meliponini) in Ubatuba, SP, Brazil.

This study describes how the foraging activity of Melipona rufiventris is influenced by the environment and/or by the state of a colony. Two colonies were studied in Ubatuba, SP (44 degrees 48 W and 23 degrees 22 S) from July/2000 to June/2001. These colonies were classified as strong (Colony 1) and intermediate (Colony 2) according to their general conditions: population and brood comb size and number of food pots. The bees were active from dawn to dusk. The number of pollen loads presented a positive correlation with relative humidity (r(s) = 0.401; p <0.01) and was highest between 70 and 90%. However, it was negatively correlated with temperature (r(s) = -0.228; p <0.01) showing a peak between 18 and 23 degrees C. The number of nectar loads presented a positive correlation with temperature (r(s) = 0.244; p <0.01) and light intensity (r(s) = 0.414; p <0.01); it was greater between 50 and 90% of relative humidity and 20 and 30 degrees C of temperature. They collected more nectar than pollen throughout the day, and were more active between 6 and 9 hours. Workers from Colony 1 (strong) collected nectar in greater amounts and earlier than those from Colony 2 (intermediate). The number of pollen, nectar and resin loads varied considerably between the study days. Peaks of pollen collection occurred earlier in months with longer days and in a hotter and more humid climate. The foraging behavior of M. rufiventris is probably affected by the state of the colony and by environmental conditions, notably temperature, relative humidity, light intensity and length of the day.     

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.     

Reproductive biology of Pachira aquatica Aubl. (Malvaceae: Bombacoideae): a tropical tree pollinated by bats,sphingid moths and honey bees

We investigated the reproductive biology, including the floral biology, pollination biology, breeding system and reproductive success, of Pachira aquatica, a native and dominant tropical tree of fresh water wetlands, throughout the coastal plain of the Gulf of Mexico. The flowers present nocturnal anthesis, copious nectar production and sugar concentration (range 18–23%) suitable for nocturnal visitors such as bats and sphingid moths. The main nocturnal visitors were bats and sphingid moths while bees were the main diurnal visitors. There were no differences in legitimate visitation rates among bats, moths and honey bees. Bats and honey bees fed mainly on pollen while moths fed on nectar, suggesting resource partitioning. Eight species of bats carried pollen but Leptonycteris yerbabuenae is probably the most effective pollinator due to its higher pollen loads. The sphingid moths Manduca rustica, Cocytius duponchel and Eumorpha satellitia were recorded visiting flowers. Hand pollination experiments indicated a predominant outcrossing breeding system. Open pollination experiments resulted in a null fruit set, indicating pollen limitation; however, mean reproductive success, according to a seasonal census, was 17 ± 3%; these contrasting results could be explained by the seasonal availability of pollinators. We conclude that P. aquatica is an outcrossing species with a pollination system originally specialized for bats and sphingid moths, which could be driven to a multimodal pollination system due to the introduction of honey bees to tropical America.     

Nectar and Pollen Sources for Honeybee （Apis cerana cerana Fabr.） in Qinglan Mangrove Area, Hainan Island, China

In the present study, nectar and pollen sources for honeybee （Apls cerana cerana Fabr.） were studied in Qlnglan mangrove area, Hainan Island, China, based on microscopic analysis of honey and pollen load （corblcular and gut contents） from honeybees collected In October and November 2004. Qualitative and quantitative melittopalynologlcal analysis of the natural honey sample showed that the honey is of unlfloral type with Mimosa pudlca L. （Mlmosaceae） as the predominant （89.14%） source of nectar and pollen for A. cerana cerana In October. Members of Araceae are an Important minor （3%-15%） pollen type, whereas those of Arecaceae are a minor （〈3%） pollen type. Pollen grains of Nypa fruticans Wurmb., Rhlzophora spp., Excoecarla agallocha L., Lumnitzera spp., Brugulera spp., Kandella candel Druce, and Ceriops tagal （Perr.） C. B. Rob. are among the notable mangrove texa growing In Qinglan mangrove area recorded as minor taxa In the honey. The absolute pollen count （I.e. the number of pollen grains/10 g honey sample） suggests that the honey belongs to Group V （〉1 000 000）. Pollen analysis from the corblcular and gut contents of A. cerana cerana revealed the highest representation （95.60%） of members of Sonneratia spp. （Sonneratlaceae）, followed by Bruguiera spp. （Rhizophoraceae）, Euphorblaceae, Poaceae, Fabaceae, Arecaceae, Araceae, Anacardlaceae, and Rublaceae. Of these plants, those belonging to Sonneratla plants are the most Important nectar and pollen sources for A. cerana cerana and are frequently foraged and pollinated by these bees in November.     

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.     

Nesting ecology and floral resource of Xylocopa augusti Lepeletier de Saint Fargeau (Hymenoptera,Apidae) in Argentina

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