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Stingless bees are important pollinators in the tropics. The tremendous variation in body size makes them an excellent group to study how miniaturization affects vision and visual behaviours. Using direct measurements and micro-CT, we reconstructed the eye structure, estimated anatomical spatial resolution and optical sensitivity of the stingless bee Tetragonula iridipennis. T. iridipennis is similar in size to the Australian stingless bee Tetragonula carbonaria and is smaller than honeybees. It has correspondingly small eyes (area = 0.56 mm 2 ), few ommatidia (2451 ± 127), large inter-facet (3.0 ± 0.6°) and acceptance angles (2.8°). Theoretical estimates suggest that T. iridipennis has poorer spatial resolution (0.17 cycles degree -1 ) than honeybees, bumblebees, and T. carbonaria. Its optical sensitivity (0.08 µm 2 sr), though higher than expected, is within the range of diurnal bees. This may provide them with greater contrast sensitivity, which is likely more relevant than the absolute sensitivity in this diurnal bee. Behaviourally determined detection thresholds for single targets using y-maze experiments were 11.5° for targets that provide chromatic contrast alone and 9.1° for targets providing chromatic and achromatic contrast. Further studies into microhabitat preferences and behaviour are required to understand how miniaturization influences its visual ecology., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

To understand how insect pollinators find flowers against complex backgrounds in diverse natural habitats, it is required to accurately estimate the thresholds for target detection. Detection thresholds for single targets vary between bee species and have been estimated in the Western honeybee, a species of bumblebee and in a stingless bee species. We estimated the angular range of detection for coloured targets in the Asian honeybee Apis cerana. Using a Y-maze experimental set up, we show that targets that provided both chromatic and green receptor contrast were detected at a minimum visual angle of 7.7°, while targets with only chromatic contrast were detected at a minimum angle of 13.2°. Our results thus provide a robust foundation for future studies on the visual ecology of bees in a comparative interspecific framework.

Fruit-feeding pteropodid bats roost under varying light conditions. Some roost in trees with high exposure to daylight (> 1000 lx), while others roost in dark caves (< 0.1 lx). To understand the effect of ambient light intensity and moon phase on flight activity, we examined flight times across five lunar cycles in three pteropodid species whose roosts differ in daylight exposure. We found significant interspecific differences in flight emergence and termination times. All species initiated flights after sunset but Rousettus leschenaultii, which typically roosts in caves, delayed emergence (40 ± 11 min) more than the two tree-roosting species Pteropus giganteus (16 ± 6 min) and Cynopterus sphinx (19 ± 7 min). R. leschenaultii terminated flights earlier (30 ± 7 min before sunrise) than P. giganteus (11 ± 11 min) and C. sphinx (16 ± 10 min). All individuals from P. giganteus and C. sphinx roosts emerged within less than an hour, while emergence times were more spread out in the R. leschenaultii colony. Peak emergence times differed across moon phases in the cave-roosting R. leschenaultii but not in the other species. Flight activity in R. leschenaultii is restricted to comparatively lower light levels than the tree-roosting species. The observed interspecific differences suggest that bat species, sharing same landscapes may respond differently to light pollution.

In addition to sugars, nectar contains multiple nutrient compounds in varying concentrations, yet little is known of their effect on the reward properties of nectar and the resulting implications for insect behaviour. We examined the pre-ingestive responses of honeybees to sucrose solutions containing a mix of pollen compounds, the amino acids proline or phenylalanine, or known distasteful substances, quinine and salt. We predicted that in taste and learning assays, bees would respond positively to the presence of nutrient compounds in a sucrose solution. However, bees' proboscis extension responses decreased when their antennae were stimulated with pollen- or amino acid-supplemented sucrose solutions. Compared to pure sucrose, bees exhibited worse acquisition when conditioned to an odour with pollen-supplemented sucrose as the unconditioned stimulus. Such learning impairment was also observed with quinine-containing sucrose solutions. Our results suggest that bees can use their antennae to detect pollen compounds in floral nectars. Depending on the type and concentrations of compounds present, this may result in nectar being perceived as distasteful by bees, making it less effective in reinforcing the learning of floral cues. Such reward devaluation might be adaptive in cases where plants benefit from regulating the frequency of bee visitation.

Large carpenter bees are charismatic and ubiquitous flower visitors in the tropics and sub-tropics. Unlike honeybees and bumblebees that have been popular subjects of extensive studies on their neuroethology, behaviour and ecology, carpenter bees have received little attention. This review integrates what is known about their foraging behaviour as well as sensory, physiological and cognitive adaptations and is motivated by their versatility as flower visitors and pollinators. This is evident from their extremely generalist foraging and adeptness at handling diverse flower types as legitimate pollinators and as illegitimate nectar robbers. They purportedly use traplining to forage between isolated patches and are long-distance flyers over several kilometres suggesting well-developed spatial learning, route memory and navigational capabilities. They have a broad range of temperature tolerance and thermoregulatory capabilities which are likely employed in their forays into crepuscular and nocturnal time periods. Such temporal extensions into dim-light periods invoke a suite of visual adaptations in their apposition optics. Thus, we propose that carpenter bees are an excellent though understudied group for exploring the complex nature of plant-pollinator mutualisms from ecological and mechanistic perspectives.

The proboscis extension conditioning (PER) is a successful behavioural paradigm for studying sensory and learning mechanisms in bees. Whilst mainly used with olfactory and tactile stimuli, more recently reliable PER conditioning has been achieved with visual stimuli such as colours and looming stripes. However, the results reported in different studies vary quite strongly, and it remains controversially discussed how to best condition visual PER. It is particularly striking that visual PER leads to more limited performance as compared to visual conditioning of free-flying bees. It could be that visual PER learning is affected by the lack of movement and that the presence of visual motion cues could compensate for it. We tested whether bees would show differences in learning performances when conditioned either with a colour and motion stimulus in combination or with colour alone. Colour acquisition was improved in the presence of the motion stimulus. The result is consistent with the idea that visual learning might be tightly linked to movement in bees, given that they use vision predominantly during flight. Our results further confirm recent findings that successful visual PER conditioning in bees is achievable without obligatorily removing the antennae.

Most bees are diurnal, with behaviour that is largely visually mediated, but several groups have made evolutionary shifts to nocturnality, despite having apposition compound eyes unsuited to vision in dim light. We compared the anatomy and optics of the apposition eyes and the ocelli of the nocturnal carpenter bee, Xylocopa tranquebarica, with two sympatric species, the strictly diurnal X. leucothorax and the occasionally crepuscular X. tenuiscapa. The ocelli of the nocturnal X. tranquebarica are unusually large (diameter ca. 1 mm) and poorly focussed. Moreover, their apposition eyes show specific visual adaptations for vision in dim light, including large size, large facets and very wide rhabdoms, which together make these eyes 9 times more sensitive than those of X. tenuiscapa and 27 times more sensitive than those of X. leucothorax. These differences in optical sensitivity are surprisingly small considering that X. tranquebarica can fly on moonless nights when background luminance is as low as 10(-5) cd m(-2), implying that this bee must employ additional visual strategies to forage and find its way back to the nest. These strategies may include photoreceptors with longer integration times and higher contrast gains as well as higher neural summation mechanisms for increasing visual reliability in dim light.

Bees are mostly active during the daytime, but nocturnality has been reported in some bee families. We studied temporal flight activity in three species of carpenter bees (genus Xylocopa) in relation to light intensities. X. leucothorax is diurnal, X. tenuiscapa is largely diurnal being only occasionally crepuscular, while X. tranquebarica is truly nocturnal. Occasional forays into dim light by X. tenuiscapa are likely to be due to the availability of richly rewarding Heterophragma quadriloculare (Bignoniaceae) flowers, which open at night. X. tranquebarica can fly even during the moonless parts of nights when light intensities were lower than 10(-5) cd m(-2), which makes this species the only truly nocturnal bee known so far. Other known dim-light species fly during crepuscular or moonlit periods. We compare eye and body sizes with other known diurnal and dim-light bees. We conclude that while extremely large ocellar diameters, large eye size:body size ratio, large number of ommatidia and large ommatidial diameters are all adaptations to dim-light foraging, these alone do not sufficiently explain the flights of X. tranquebarica in extremely dim light. We hypothesise that additional adaptations must confer extreme nocturnality in X. tranquebarica.