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Pollen Use in Bees and Other Insects
Session leader: Margrit McIntosh.
Main sources for this introduction: Burgess (1991), Gess (1996), Kevan and Baker (1983), Labandeira (1997)
Pollen use by bees (Hymenoptera: Apoidea) is described in detail in the up-to-date and comprehensive review by Wcislo and Cane (1996); hence, in this introduction I will focus on pollen use in other insects.
Pollen grains consist of a nutrient-rich inner protoplasm, surrounded by an exceptionably durable coat ("exine"), and the grains are often covered by oil-rich and fragrance-rich "pollenkit" (a substance derived from tapetal tissue in the anther). Pollen contains abundant protein and free amino acids, although the proportion of these nutrients can vary widely among plant taxa. The protein is intended for the growth of the pollen tube, and it is this nutrient in particular that makes pollen an appropriate food for the developmental stages of insects. Lipids are always present;
starch may or may not be present.
There is abundant evidence for pollen- and spore-eating by insects in the fossil record. Pollen evolved from the microspores of seedless vascular plants, appearing in the fossil record during the Carboniferous. Spores and pollen ingested and excreted by insects during the Late Carboniferous to Miocene eras are sometimes even identifiable to source plant taxa (Labandeira 1997). Fossil insects associated with pollen feeding include thysanurans, diaphanopterodean nymphs, adult hemipteroids, protorthopterans, and grylloblattids (Labandeira 1997). Modern pollen-eating insects occur in at least 14 orders (Table 1).
Insects have evolved several different methods for ingesting pollen grain, and the mode of ingestion may or may not be associated with specialized mouthparts. The most common modes include:
- Swallowing the grains whole.
Following ingestion, the grains may burst due to osmotic pressure. Or the contents may be digested and absorbed through the pores in the exine of the grains.
- Chewing up the grains.
Pollen exine fragments are not digested, and may be identified in insect excretions.
=> Bee larvae either chew or swallow pollen grains.
- The punch-and-suck method (e.g., certain thrips)
The insect mouthparts are used to pierce the grain through one of the pores, and the contents are extracted without swallowing the grains.
There appear to be two primary paths by which pollen-feeding has been arrived at, evolutionarily, by insects. One is via carnivory, and in particular, feeding on other insects. This seems to be the path taken by bees, who share a common ancestor with the exclusively carnivorous sphecid wasps; by masarine wasps in the family Vespidae; and by (e.g.) staphylinid beetles. Because it is a rich source of protein, pollen is nutritionally much more similar to insects themselves than it is to non-reproductive plant parts. The second path is that taken by phytophagous insects which come to feed on the flowers of plants, instead of on foliar tissue or other parts. Some insects are general florivores, while others are specialized to pollen or ovules.
A particularly interesting case is that of the masarine wasps (Hymenoptera: Vespidae: Masarinae), which have independently evolved the habit of provisioning their larvae with a mixture of pollen and nectar, as do bees. Some masarines carry the pollen in their crop and then regurgitate it to form the provisions, as do some bees (Colletidae: Hylaeinae). Others have developed brushes on their legs for carrying pollen, such as those possessed by bees -- but these masarine wasps have them on the forelegs, instead of the hindlegs! In many cases it is thought that masarine wasps are effective pollinators of the plants they visit. I find it particularly fascinating that most masarines seems to be specialists on their pollen hosts, as are most bees (and, of course, as are most herbivorous insects). There are only about 300 known species of masarines, compared to the >20,000 spp. of bees.
Just in the past year, a report has been published of a pollen-collecting sphecid wasp in Sri Lanka (Krombein & Norden 1997; Hymenoptera: Sphecidae: Crabroninae), so it would appear that the habit of collecting pollen instead of insects to provision the larvae has evolved three separate times in the Hymenoptera.
Table 1: Insects that eat pollen. Compiled from Burgess (1991), and Kevan & Baker (1983)
COLLEMBOLA
|
BLATTARIA
|
DERMAPTERA |
ORTHOPTERA |
PLECOPTERA |
HEMIPTERA
|
Anthocoridae
Miridae
|
THYSANOPTERA |
NEUROPTERA |
MECOPTERA |
COLEOPTERA
|
Alleculidae
Buprestidae
Cantharidae
Cephaloidae
Cerambycidae
Cleridae
Curculionidae
Dermestidae
Languriidae
Meloidae
Melyridae
Mordellidae
Nitidulidae
Oedemeridae
Phalacridae
Scarabidae |
DIPTERA
|
Anthomyiidae
Bibionidae
Bombyliidae
Calliphoridae
Mycetophilidae
Muscidae
Scatopsidae
Syrphidae
Tachinidae |
LEPIDOPTERA
|
Micropterygidae
Nyphalidae |
TRICHOPTERA |
HYMENOPTERA
|
Xyelidae
Apoidea
Vespidae (Masarinae)
Sphecidae (one species; Krombein & Norden 1997)
|
Required papers
- Wcislo, W.T., and J.H. Cane. 1996. Floral resource utilization by solitary bees (Hymenoptera: Apoidea) and exploitation of their stored foods by natural enemies. Annual Review of Entomology 41:257-286.
- Muller, A. 1996. Host-plant specialization in western Palearctic Anthidiine bees (Hymenoptera: Apoidea: Megachilidae). Ecological Monographs 66(2):235-257.
Full references list
General References
- Baker, H.G., and I. Baker. 1979. Starch in angiosperm pollen grains and its evolutionary significance. American Journal of Botany 66:591-600.
- Boucot, A. J. 1990. Evolutionary palaeobiology of behavior and coevolution. Amsterdam: Elsevier.
- Burgess, K.H. 1991. Florivory: the ecology of flower feeding insects and their host plants. Ph.D. dissertation, Harvard University, Cambridge, MA.
- Kato, M., and T. Inoue. 1994. Origin of insect pollination. Nature 368:195.
- Kato, M., T. Inoue, and T. Nagamitsu. 1995. Pollination biology of Gnetum (Gnetaceae) in a lowland mixed dipterocarp forest in Sarawak. American Journal of Botany 82:862-868.
- Kevan, P. G., and H. G. Baker. 1983. Insects as flower visitors and pollinators. Annual Review of Entomology 28:407-453.
- Labandeira, C. C. 1997. Insect mouthparts: ascertaining the paloebiology of insect feeding strategies. Annual Review of Ecology and Systematics 28:153-193.
- Pellmyr, O. 1992. Evolution of insect pollination and angiosperm diversification. Trends in Ecology & Evolution 7:46-49.
- Proctor, M., P. Yeo, and A. Lack. 1996. The natural history of pollination. Portland, Oregon, USA: Timber Press.
- Taylor, T. N., and A. C. Scott. 1987. Interactions of plants and animals during the Carboniferous. BioScience 33 (8):488-493.
- Vogel, S. 1978. Evolutionary shifts from reward to deception in pollen flowers. In The pollination of flowers by insects, edited by A. J. Richards. London: The Academic Press.
Pollen Use in Bees
- Danforth, B. N. 1994. Taxonomic review of Calliopsis subgenus Hypomacrotera (Hymenoptera: Andrenidae), with special emphasis on the distributions and host plant associations. Pan-Pacific Entomologist 70 :283-300.
- Dobson, H. E. M. 1987. Role of flower and pollen aromas in host-plant recognition by solitary bees. Oecologia 72:618-623.
- Dobson, H. E. M. 1988. Survey of pollen and pollenkitt lipids: chemical cues to flower visitors? American Journal of Botany 75:170-182.
- Dobson, H. E. M., I. Groth, and G. Bergstrom. 1996. Pollen advertisement: chemical contrasts between whole-flower and pollen odors. American Journal of Botany 83 :877-885.
- Dobson, H. E. M., and Y.-s. Peng. 1997. Digestion of pollen components by larvae of the flower-specialist bee Chelostoma florisomne (Hymenoptera: Megachilidae). Journal of Insect Physiology 43 (1):89-100.
- Eickwort, G. C., and H. S. Ginsberg. 1980. Foraging and mating behavior in Apoidea. Annual Review of Entomology 25:421-446.
- Harder, L. D., and R. M. R. Barclay. 1994. The functional significance of poricidal anthers and buzz pollination: controlled pollen removal from Dodecatheon. Functional Ecology 8:509-517.
- Linsley, E. G., and J. W. MacSwain. 1958. The significance of floral constancy among bees of the genus Diadasia (Hymenoptera: Anthophoridae). Evolution 12 :219-223.
- Minckley, R. L., W. T. Wcislo, D. Yanega, and S. L. Buchmann. 1994. Behavior and phenology of a specialist bee (Dieunomia) and sunflower (Helianthus) pollen availability. Ecology 75:1406-1419.
- Muller, A. 1996. Convergent evolution of morphological specializations in Central European bee and honey wasp species as an adaptation to the uptake of pollen from nototribic flowers (Hymenoptera: Apoidea and Masaridae). Biological Journal of the Linnean Society 57 (3):235-252.
- Muller, A. 1995. Morphological specializations in Central European bees for the uptake of pollen from flowers with anthers hidden in narrow corolla tubes (Hymenoptera, Apoidea). Entomologia Generalis 20 (1-2):43-57.
- Roubik, D. W. 1989. Ecology and natural history of tropical bees. Cambridge: Cambridge University Press.
- Strickler, K. 1979. Specialization and foraging efficiency of solitary bees. Ecology 60:998-1009.
- Thorp, R. W. 1979. Structural, behavioral, and physiological adaptations of bees (Apoidea) for collecting pollen. Annals of the Missouri Botanical Garden 66:788-812.
- Westerkamp, C. 1996. Pollen in bee-flower relations: some considerations on melittophily. Botanica Acta 109:325-332.
- Westerkamp, C. 1997. Keel blossoms: bee flowers with adaptations against bees. Flora 192 (2):125-132.
Pollen Use in Masarine Wasps
- Gess, S. K. 1992. Biogeography of the Masarine wasps (Hymenoptera, Vespidae, Masarinae), with particular emphasis on the southern African taxa and on correlations between Masarine and forage plant distributions. Journal of Biogeography 19:491-503.
- Gess, S. K. 1996. The pollen wasps: ecology and natural history of the Masarinae. Cambridge, Massachusetts, USA: Harvard University Press.
- Houston, T. F. 1984. Bionomics of a pollen-collecting wasp, Paragia tricolor (Hymenoptera: Vespidae: Masarinae), in Western Australia. Rec. West. Aust. Mus. 11:141-151.
- Muller, A. 1996. Convergent evolution of morphological specializations in Central European bee and honey wasp species as an adaptation to the uptake of pollen from nototribic flowers (Hymenoptera: Apoidea and Masaridae). Biological Journal of the Linnean Society 57 (3):235-252.
- Neff, J. L., and B. B. Simpson. 1985. Hooked setae and narrow tubes: foretarsal pollen collection by Trimeria buyssoni (Hymenoptera: Masaridae). Journal of the Kansas Entomological Society 58:730-732.
- Torchio, P. F. 1970. The ethology of the wasp Pseudomasaris edwardsii (Cresson), and a description of its immature forms (Hymenoptera: Vespoidea, Masaridae). Pp. 32 in Los Angeles County Musuem Contributions in Science: Los Angeles County Museum.
- Torchio, P. F. 1974. Mechanisms involved in the pollination of Penstemon visited by the masarid wasp Pseudomasaris vespoides (Cresson) (Hymenoptera: Vespoidea). Pan-Pacific Entomologist 50:226-234.
Pollen Use in Other Insects
- Charpentier, R. 1985. Host plant selection by the pollen beetle Meligethes aeneus. Entomologia Experimentalis et Applicata 38:277-285.
- Ekbom, B., and A. Borg. 1996. Pollen beetle (Meligethes aeneus) oviposition and feeding preference on different host plant species. Entomologia Experimentalis et Applicata 78 (3):291-299.
- Evans, K. A., and L. J. Allenwilliams. 1994. Laboratory and field response of the pollen beetle, Meligethes aeneus, to the odour of oilseed rape. Physiological Entomology 19 (4):285-290.
- Giamoustaris, A., and R. Mithen. 1996. The effect of flower colour and glucosinolates on the interaction between oilseed rape and pollen beetles. Entomologia Experimentalis et Applicata 80:206-208.
- Grant, V., and W. A. Connell. 1979. The association between Carpophilus beetles and cactus flowers. Plant Systematics and Evolution 133:99-102.
- Hickman, J.M.; Lovei, G.L., and Wratten, S.D. 1995. Pollen feeding by adults of the hoverfly Melanostoma fasciatum (Diptera, Syrphidae). New Zealand Journal of Zoology 22(4):387-392.
- Hunt, J. H., P. W. Brown, K. M. Sago, and J. A. Kerker. 1991. Vespid wasps eat pollen (Hymenoptera: Vespidae). Journal of the Kansas Entomological Society 64 (2):127-130.
- Kirk, W. D. J. 1985. Pollen-feeding and the host specificity and fecundity of flower thrips (Thysanoptera). Ecological Entomology 10:281-289.
- Krombein, K.V., and Norden, B.B. 1997. Nesting behavior of Krombeinictus nordenae Leclercq, a sphecid wasp with vegetarian larvae (Hymenoptera, Sphecidae, Crabroninae). Proceedings of the Entomological Society of Washington 99(1):42-49.
- Norstog, K. J., D. W. Stevenson, and K. J. Niklas. 1986. The role of beetles in the pollination of Zamia furfuracea L. fil. (Zamiaceae). Biotropica 18:300-306.
- Wacht, S., K. Lunau, and K. Hansen. 1996. Optical and chemical stimuli control pollen feeding in the hoverfly Eristalis tenax. Entomologia Experimentalis et Applicata 80:50-53.
Copyright 1998, Margrit Mcintosh.