Morphologically complex flowers are characterized by bilateral symmetry, tube-like shapes, deep corolla tubes, fused petals, and/or poricidal anthers, all of which constrain the access of insect visitors to floral nectar and pollen rewards. Only a subset of potential pollinators, mainly large bees, learn to successfully forage on such flowers. Thus, complexity may comprise a morphological filter that restricts the range of visitors and thereby increases food intake for successful foragers. Such pollinator specialization, in turn, promotes flower constancy and reduces cross-species pollen transfer, providing fitness benefits to plants with complex flowers. Since visual signals associated with floral morphological complexity are generally honest (i.e., indicate food rewards), pollinators need to perceive and process them. Physiological studies show that bees detect distant flowers through long-wavelength sensitive photoreceptors. Bees effectively perceive complex shapes and learn the positions of contours based on their spatial frequencies. Complex flowers require long handling times by naive visitors, and become highly profitable only for experienced foragers. To explore possible pathways towards the evolution of floral complexity, we discuss cognitive mechanisms that potentially allow insects to persist on complex flowers despite low initial foraging gains, suggest experiments to test these mechanisms, and speculate on their adaptive value.
Bees exemplify flights under bright sunlight. A few species across bee families have evolved nocturnality, displaying remarkable adaptations to overcome limitations of their daylight-suited apposition eyes. Phase inversion to nocturnality in a minority of bees that co-exist with diurnal bees provides a unique opportunity to study ecological benefits that mediate total temporal niche shifts. While floral traits and sensory modalities associated with the evolution of classical nocturnal pollination syndromes, e.g. by bats and moths, are well-studied, nocturnality in bees represents a poorly understood, recently invaded, extreme niche. To test the competitive release hypothesis, we examine how nocturnality shapes foraging by comparing pollen loads, nest pollen, and flower visitation of sympatric nocturnal and diurnal carpenter bees. We predicted that nocturnal bees primarily use night-blooming flowers, show little/no resource overlap with diurnal species and competitive release favors night-time pollen collection for provisioning. Contrarily, we found substantial resource overlap between nocturnal and diurnal bees. Flower opening times, floral longevity and plant abundance did not define nocturnal flower use. Smaller pollen loads on nocturnal foragers suggest subsistence on resource leftovers largely from diurnal flowers. Greater pollen types/diversity on nocturnal foragers indicate lower floral constancy compared to diurnal congenerics. Reduced activity during new moon compared to full moon suggests constraints to nocturnal foraging. Invasion and sustenance within the nocturnal niche is characterized by: (i) opportunistic foraging on residual resources as indicated by smaller pollen loads, extensive utilization of day-blooming flowers and substantial overlap with diurnal bees, (ii) generalization at two levels-between and within foraging trips as indicated by lower floral constancy, (iii) reduced foraging on darker nights, indicating visual constraints despite sensitive optics. This together with smaller populations and univoltine breeding in nocturnal compared to multivoltine diurnal counterparts suggest that nocturnality imposes substantial fitness costs. In conclusion, the evolution of nocturnality in bees is accompanied by resource generalization instead of specialization. Reduced floral constancy suggests differences in foraging strategies of nocturnal and diurnal bees which merits further investigation. The relative roles of competition, floral rewards and predators should be examined to fully understand the evolution and maintenance of nocturnality in bees.
The importance of pollinators as selective agents for many floral traits is well established, but understanding their role in the evolution of complex floral shapes remains challenging. This is because pollinators often need much practice to efficiently handle morphologically complex flowers and extract their food rewards. What induces foragers to persistently visit and pollinate complex flowers despite their initial low profitability? We previously found that naive bumblebees, and unsuccessful feeding attempts of experienced ones, contribute to the pollination of complex flowers. Here we tested a complementary hypothesis, positing that successful foraging on flowers of one complex shape prepares pollinators to visit other species of different complex morphologies. We trained bumblebees to computer-controlled artificial flowers that were either simple, complex or both simple and complex. We then recorded their feeding choices and handling times on a second array of simple and complex flowers that had different shapes and required another handling technique. Bees trained on a single flower type (whether simple or complex) preferred flowers of the same type in the testing array. The foragers’ preferences after training on both flower types depended on the reward schedule during training: when both flower types rewarded equally, simple flowers were preferred at the test phase; when complex flowers provided higher reward during training, they became the preferred flower type during testing. These results suggest that successful foraging on complex flowers, especially when highly rewarding, can indeed induce insect pollinators to attempt additional flower species with other complex shapes.
Little is known about the role of crabs as seed dispersers and predators. Recently, there has been interest in understanding their influence on plant recruitment in coastal forests. Secondary seed removal by crabs in a swampspecialist tree, Myristica fatua, was investigated in the rare and patchy freshwater Myristica swamps in the Western Ghats in India. Tethered-line experiments were used to determine the role of crabs as secondary seed-removal agents in two study sites. Crabs transported a large percentage (63.3%) of seeds (n = 60) placed on the forest floor compared with rodents (25%) and other unknown agents (13.3%). Simultaneous choice experiments suggested that the nutrient-rich arils covering seeds were consumed, but there was no evidence for seed predation by crabs. A small percentage (13.3%) of monitored seeds (n = 60) germinated from within crab burrows. The spatial scale of secondary removal by crabs was restricted to < 10 m. In these fragmented swamp forests, secondary removal by crabs retains seeds largely within the swamps, where conditions for their establishment and survival are optimal. Thus, secondary seed removal by crabs could provide temporal and spatial refugia from seed predators such as rodents in Myristica fatua.
Social insects have evolved a variety of architectural formations. Bees and wasps are well known for their ability to achieve compact structures by building hexagonal cells. Polistes wattii, an open nesting paper wasp species, builds planar hexagonal structures. Here, using the pair correlation function approach, we show that their nests exhibit short-range hexagonal order (no long-range order) akin to amorphous materials. Hexagonal orientational order was well preserved globally. We also show the presence of topological defects such as dislocations (pentagon-heptagon disclination pairs) and Stone-Wales quadrupoles, and discuss how these defects were organised in the nest, thereby restoring order. Furthermore, we suggest the possible role of such defects in shaping nesting architectures of other social insect species.
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