Hostname: page-component-cd9895bd7-p9bg8 Total loading time: 0 Render date: 2024-12-25T05:49:19.572Z Has data issue: false hasContentIssue false

Flies and flowers: taxonomic diversity of anthophiles and pollinators

Published online by Cambridge University Press:  31 May 2012

B.M.H. Larson
Affiliation:
Department of Ecology, Evolution and Marine Biology, University of California – Santa Barbara, Santa Barbara, California, United States of America 93106
P.G. Kevan*
Affiliation:
Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada N1G 2W1
D.W. Inouye
Affiliation:
Department of Zoology, University of Maryland, College Park, Maryland, United States of America 20742
*
1 Author to whom all correspondence should be addressed (E-mail: [email protected]).

Abstract

The Diptera are the second most important order among flower-visiting (anthophilous) and flower-pollinating insects worldwide. Their taxonomic diversity ranges from Nematocera to Brachycera, including most families within the suborders. Especially important are Syrphidae, Bombyliidae, and Muscoidea. Other families, especially of small flies, are less appreciated and often overlooked for their associations with flowers. We have compiled records of their flower visitations to show that they may be more prevalent than usually thought. Our knowledge of anthophilous Diptera needs to be enhanced by future research concerning (i) the significance of nocturnal Nematocera and acalypterate muscoids as pollinators, (ii) the extent to which the relatively ineffective pollen-carrying ability of some taxa can be compensated by the abundance of individuals, and (iii) the role of Diptera as pollinators of the first flowering plants (Angiospermae) by using phylogenetic and palaeontological evidence. Specializations in floral relationships involve the morphology of Diptera, especially of their mouthparts, nutritional requirements, and behaviour, as well as concomitant floral attributes. The South African flora has the most highly specialized relations with dipterous pollinators, but in arctic and alpine generalist fly–flower relations are important in pollination and fly nutrition.

Résumé

Les diptères occupent le second rang au monde parmi les ordres d’insectes qui visitent les fleurs (anthophiles) et qui assurent la pollinisation. Leur diversité taxonomique va des nématocères aux brachycères et inclut presque toutes les familles de ces sous-ordres, en particulier les Syrphidae, les Bombyliidae et les Muscoidea qui occupent une place prépondérante. Les associations des autres familles avec les fleurs, particulièrement les familles de mouches de petite taille, passent souvent inaperçues. Nous avons compilé des données sur leurs visites aux fleurs dans le but de démontrer qu’elles sont probablement plus importantes qu’on ne le croit. Nos connaissances des diptères anthophiles doivent être complétées par d’autres recherches (i) pour déterminer l’importance des nématocères nocturnes et des muscoïdes acalyptères comme pollinisateurs, (ii) pour évaluer jusqu’à quel point l’inefficacité relative de certains taxons comme transporteurs de pollen peut être compensée par l’abondance des individus et (iii) pour juger du rôle des diptères comme pollinisateurs des premières plantes à fleurs (Angiospermae) en utilisant des données phylogénétiques et paléontologiques. Les spécialisations de relations fleurs–insectes sont basées sur des particularités morphologiques des diptères, surtout de leurs pièces buccales, sur leurs besoins nutritifs et sur leur comportement, en même temps que sur les attributs correspondants des fleurs. La flore de l’Afrique du Sud est celle qui a les relations les plus spécialisées avec les diptères pollinisateurs. Cependant, dans les régions arctiques et alpines, des relations apparemment non spécialisées entre les mouches et les fleurs sont importantes pour la pollinisation des fleurs et l’alimentation des mouches.

[Traduit par la Rédaction]

Type
Articles
Copyright
Copyright © Entomological Society of Canada 2001

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Ackerman, J.D., Mesler, M.R. 1979. Pollination biology of Listera cordata (Orchidaceae). American Journal of Botany 66: 820–4CrossRefGoogle Scholar
Addicott, J.F. 1986. Variation in the costs and benefits of mutualism: the interaction between yuccas and yucca moths. Oecologia 70: 486–94CrossRefGoogle ScholarPubMed
Agnew, J.D. 1976. A case of myophily involving Drosophilidae (Diptera). Journal of South African Botany 42: 8595Google Scholar
Ambrose, J.D., Kevan, P.G., Gadawski, R.M. 1985. Hop tree (Ptelea trifoliata) in Canada: population and reproductive biology of a rare species. Canadian Journal of Botany 63: 1928–35CrossRefGoogle Scholar
Arroyo, M.T.K., Primack, R., Armesto, J. 1982. Community studies in pollination ecology in the high temperate Andes of central Chile. I. Pollination mechanisms and altitudinal variation. American Journal of Botany 69: 8297CrossRefGoogle Scholar
Arruda VLV de-, Sazima 1996. Flores vistadas por sirfideos (Diptera: Syrphidae) en mata mesofila de Campinas, SP. Revista Brasileira de Botanica 19: 109–17Google Scholar
Atwood, J.T. 1982. How is Paphiopedilum pollinated? American Orchid Society Bulletin 51: 1057–8Google Scholar
Bänziger, H. 1991. Stench and fragrance: unique pollination lure of Thailand's largest flower, Rafflesia kerrii Meijer. Natural History Bulletin of the Siam Society 39: 1952Google Scholar
Barrett, S.C.H., Helenurm, K. 1987. The reproductive biology of boreal forest herbs. I. Breeding systems and pollination. Canadian Journal of Botany 65: 2036–46CrossRefGoogle Scholar
Barth, F.G. 1985. Insects and flowers. The biology of a partnership. Princeton: Princeton University PressGoogle Scholar
Baumann, E. 1978. Rennfliegen (Diptera: Phoridae) als Blutenbesucher. Kritische Sichtung der Literatur Flora 167: 301–14CrossRefGoogle Scholar
Beaman, R.S., Decker, P.J., Beaman, J.H. 1988. Pollination of Rafflesia (Rafflesiaceae). American Journal of Botany 75: 1148–62CrossRefGoogle Scholar
Beattie, A.J. 1971. Pollination mechanisms in Viola. New Phytologist 70: 343–60CrossRefGoogle Scholar
Beattie, A.J. 1972. Insect vistors to three species of violet (Viola) in England. Entomologists' Monthly Magazine 108: 711Google Scholar
Beattie, A.J. 1976. Plant dispersion, pollination and gene flow in Viola. Oecologia 25: 291300CrossRefGoogle ScholarPubMed
Bishop, A. 1996. Field guide to the orchids of New South Wales and Victora. Sydney: University of New South Wales PressGoogle Scholar
Borror, D.J., Triplehorn, C.A., Johnson, N.F. 1989. An introduction to the study of insects. 6th ed. Philadelphia: Saunders College PublishingGoogle Scholar
Brackley, F.E., Burger, J.F. 1980. Occurrence of two anthophilous diptera on Geum radiatum (Rosaceae) in North Carolina. Entomological News 91: 110–2Google Scholar
Brantjes, N.B.M. 1980. Flower morphology of Aristolochia species and the consequences for pollination. Acta Botanica Neerlandica 29: 212–3Google Scholar
Brantjes, N.B.M. 1981. Ant, bee and fly pollination in Epipactis palustris (L.) Crantz (Orchidaceae). Acta Botanica Neerlandica 30: 5968CrossRefGoogle Scholar
Brantjes, N.B.M., Leemans, JAAM. 1976. Silene otites (Caryophyllaceae) pollinated by nocturnal Lepidoptera and mosquitoes. Acta Botanica Neerlandica 25: 281–95CrossRefGoogle Scholar
Buchmann, S.L. 1983. Buzz pollination in Angiosperms. pp 73113in Jones, C.E., Little, R.J. (Eds), Handbook of experimental pollination biology. New York: Van Nostrand ReinholdGoogle Scholar
Bultman, T.L., White, J.F. 1988. “Pollination” of a fungus by a fly. Oecologia 75: 317–9CrossRefGoogle ScholarPubMed
Burgin, S.G., Hunter, F.F. 1997 a. Nectar versus honeydew as sources of sugar for male and female black flies (Diptera: Simuliidae). Journal of Medical Entomology 34: 605–8CrossRefGoogle ScholarPubMed
Burgin, S.G., Hunter, F.F. 1997 b. Evidence of honeydew feeding in black flies (Diptera: Simuliidae). The Canadian Entomologist 129: 859–69CrossRefGoogle Scholar
Burgin, S.G., Hunter, F.F. 1997 c. Sugar meals used by female black flies (Diptera: Simuliidae): a four-habitat study. Canadian Journal of Zoology 75: 1066–72CrossRefGoogle Scholar
Bystrak, P.G., Wirth, W.W. 1978. The North American species of Forcipomyia, subgenus Euprojoannisia (Diptera: Ceratopogonidae). US Department of Agriculture Technical Bulletin 1591Google Scholar
Cameron, R.G., Troilo, D. 1982. Fly-mediated spore dispersal in Splachnum ampullaceum (Musci). Michigan Botanist 21: 5965Google Scholar
Carson, H.L., Okada, T. 1982. Ecology and evolution of flower-breeding pomace flies of New Guinea (Diptera: Drosophilidae). Entomologia Generalis 8: 13–6CrossRefGoogle Scholar
Chase, M.W. 1985. Pollination of Pleurothallis endotrachys. American Orchid Society Bulletin 54: 431–4Google Scholar
Chernov, Y.I. 1966. Kompleks antophil'nikh nacekomikh v tundrovoy zonye. [Complex of anthophilous insects in the tundra zone.] Voprosi Geografii 69: 7697 [in Russian]Google Scholar
Christensen, D.E. 1992. Notes on the reproductive biology of Stelis argentata Lindl. (Orchidaceae: Pleurothallidinae) in eastern Ecuador. Lindleyana 7: 2833Google Scholar
Coleman, E. 1934. Pollination of Pterostylis acuminata R. Br., and Pterostylis falcata Rogers. Victorian Naturalist 50: 248–52Google Scholar
Crosswhite, F.S., Crosswhite, C.D. 1984. The southwestern pipevine (Aristolochia watsonii) in relation to snakeroot oil, swallowtail butterflies, and ceratopogonid flies. Desert Plants 6: 203–7Google Scholar
Cruden, R.W. 1972. Pollination biology of Nemophila menziesii (Hydrophyllaceae) with comments on the evolution of oligolectic bees. Evolution 26: 373–89CrossRefGoogle ScholarPubMed
Currah, L., Ockendon, D.J. 1983. Onion pollination by blowflies and honeybees in large cages. Annals of Applied Biology 103: 497506CrossRefGoogle Scholar
Currah, L., Ockendon, D.J. 1984. Pollination activity by blowflies and honeybees on onions in breeders' cages. Annals of Applied Biology 105: 167–76CrossRefGoogle Scholar
Dag, A., Eisenstein, D., Gazit, S. 1998. Effect of pollen donor distance and selective fruit abscission on out-crossing rate and yield in “Tommy Atkins” mango. Journal of the American Society of Horticultural Science 123: 618–22CrossRefGoogle Scholar
D'Arcy-Burt, S., Blackshaw, R.P. 1991. Bibionids (Diptera: Bibionidae) in agricultural land: a review of damage, benefits, natural enemies and control. Annals of Applied Biology 118: 695708CrossRefGoogle Scholar
Davies, D.M., Petersen, B.V. 1956. Observations on the mating, feeding, ovarian development, and oviposition of adult black flies (Simuliidae, Diptera). Canadian Journal of Zoology 34: 615–55CrossRefGoogle Scholar
Dethier, V.G. 1976. The hungry fly: a physiological study of the behavior associated with feeding. Cambridge: Harvard University PressGoogle Scholar
Dexter, J.S. 1913. Mosquitoes pollinating orchids. Science (Washington, DC) 37: 867CrossRefGoogle ScholarPubMed
Deyrup, M.A. 1988. Pollen-feeding in Poecilognathus punctipennis (Diptera: Bombyliidae). Florida Entomologist 71: 597605CrossRefGoogle Scholar
Disney, R.H.L. 1980. Records of flower visiting by scuttle flies (Diptera: Phoridae) in the British Isles. Naturalist 105: 4550Google Scholar
Downes, J.A. 1955. The food habits and distribution of Atrichopogon pollinivorus sp. n. (Diptera: Ceratopogonidae). Transactions of the Royal Entomological Society of London 106: 439–53CrossRefGoogle Scholar
Downes, J.A. 1958. The feeding habits of biting flies and their significance in classification. Annual Review of Entomology 3: 249–66CrossRefGoogle Scholar
Downes, J.A. 1970. The feeding and mating behaviour of the specialized Empidinae (Diptera): observations on four species of Rhamphomyia in the High Arctic and a general discussion. The Canadian Entomologist 102: 769–91CrossRefGoogle Scholar
Downes, J.A., Smith, S.M. 1969. New or little known feeding habits in Empididae (Diptera). The Canadian Entomologist 101: 404–8CrossRefGoogle Scholar
Drabble, E., Drabble, H. 1917. The syrphid visitors to certain flowers. New Phytologist 16: 105–9CrossRefGoogle Scholar
Drabble, E., Drabble, H. 1927. Some flowers and their dipteran visitors. New Phytologist 26: 115–23CrossRefGoogle Scholar
Dressler, R.L. 1981. The orchids: natural history and classification. Cambridge and London: Harvard University PressGoogle Scholar
Eberle, G. 1976. Schnabelfliegen (Rhingia) als leistungsfahige Bestaubungsvermittler hochentwickelter Nektarbluten (Diptera, Syrphidae). Entomologische Zeitschrift 86: 128–36Google Scholar
Endress, P.K. 1980. The reproductive structures and systematic position of the Austrobaileyaceae. Botanische Jahrbüecher für Systematik 101: 393433Google Scholar
Entwistle, P.F. 1972. Pests of cocoa. London: LongmanGoogle Scholar
Erhardt, A. 1993. Pollination of the edelweiss, Leontopodium alpinum. Botanical Journal of the Linnean Society 111: 229–40CrossRefGoogle Scholar
Faegri, K., van der Pijl, L. 1979. The principles of pollination ecology. 3rd ed. Oxford: Pergamon PressGoogle Scholar
Faulkner, G.J. 1962. Blowflies as pollinators of brassica crops. Commercial Grower 3457: 807–9Google Scholar
Feil, J.P. 1992. Reproductive ecology of dioecious Siparuna (Monimiaceae) in Ecuador — a case of gall midge pollination. Botanical Journal of the Linnean Society 110: 171203CrossRefGoogle Scholar
Free, J.B. 1993. Insect pollination of crops. 2nd ed. London: Academic PressGoogle Scholar
Gilbert, F.S. 1981. Foraging ecology of hover flies (Diptera: Syrphidae): morphology of the mouthparts in relation to feeding on nectar and pollen in some common urban species. Ecological Entomologist 6: 245–62CrossRefGoogle Scholar
Gilbert, F., Jervis, M. 1998. Functional, evolutionary and ecological aspects of feeding-related mouthpart specializations in parasitoid flies. Biological Journal of the Linnean Society 63: 495535CrossRefGoogle Scholar
Goldblatt, P., Bernhardt, P. 1990. Pollination biology of Nivenia (Iridaceae) and the presence of heterostylous self-compatibility. Israel Journal of Botany 39: 93111Google Scholar
Goldblatt, P., Manning, J.C. 1999. The long-proboscid fly pollination system in Gladiolus (Iridaceae). Annals of the Missouri Botanical Garden 86: 758–74CrossRefGoogle Scholar
Goldblatt, P., Manning, J.C. 2000. The long-proboscid fly pollination system in southern Africa. Annals of the Missouri Botanical Garden 87: 146–70CrossRefGoogle Scholar
Goldblatt, P., Bernhardt, P., Manning, J.C. 2000. Adaptive raiation of pollination mechanisms in Ixia (Iridaceae: Crocoidea). Annals of the Missouri Botanical Garden 87: 564–77CrossRefGoogle Scholar
Goldblatt, P., Manning, J.C., Berhardt, P. 1997. Notes on the pollination of Gladiolus brevifolius (Iridaceae) by bees (Anthophoridae) and bee mimicking flies (Psilodera: Acroceridae) Journal of Kansas Enotmological Society 70: 297304Google Scholar
Goldblatt, P., Manning, J.C., Bernhardt, P. 2001. Radiation of pollination systems in Gladiolus (Iridaceae: Ixioideae) in southern Africa. Annals of the Missouri Botanical Gardens. In pressGoogle Scholar
Gorham, J.R. 1976. Orchid pollination by Aedes mosquitoes in Alaska. American Midland Naturalist 95: 208–10CrossRefGoogle Scholar
Graenicher, S. 1910. The bee-flies (Bombyliidae) in their relations to flowers. Bulletin of the Wisconsin Natural History Society 8: 91101Google Scholar
Grant, V., Grant, K.A. 1965. Flower pollination in the Phlox family. New York and London: Columbia University PressGoogle Scholar
Grensted, L.W. 1947. Diptera in the spathes of Arum maculatum L. Entomologists' Monthly Magazine 83: 13Google Scholar
Grimaldi, D. 1988. Bee flies and bluets: Bombylius (Diptera: Bombyliidae) flower-constant on the distylous species, Hedyotis caerulea (Rubiaceae), and the manner of foraging. Journal of Natural History 22: 110CrossRefGoogle Scholar
Grimaldi, D. 1999. The co-radiations of pollinating insects and angiosperms in Cretaceous. Annals of the Missouri Botanical Garden 86: 373406CrossRefGoogle Scholar
Grimstad, P.R., DeFoliart, G.R. 1974. Nectar sources of Wisconsin mosquitoes. Journal of Medical Entomology 11: 331–41CrossRefGoogle ScholarPubMed
Grinfel'd, E.K. 1955. The feeding of hover flies (Diptera: Syrphidae) and their role in the pollination of plants. Entomologicheskoe Obozrenie 34: 164–6Google Scholar
Hall, D.W., Brown, B.V. 1993. Pollination of Aristolochia littoralis (Aristolochiales: Aristolochiaceae) by males of Megaselia spp. (Diptera: Phoridae). Annals of the Entomological Society of America 86: 609–13CrossRefGoogle Scholar
Haslett, J.R. 1989. Interpreting patterns of resource utilization: randomness and selectivity in pollen feeding by adult hover flies. Oecologia 78: 433–42CrossRefGoogle Scholar
Haslett, J.R., Entwistle, P.F. 1980. Further notes on Eriozona syrphoides (Fall.) (Diptera, Syrphidae) in Hafren Forest, mid-Wales. Entomologists' Monthly Magazine 116: 36Google Scholar
Heard, T.A., Vithanage, V., Chacko, E.K. 1990. Pollination biology of cashew in the Northern Territory of Australia. Australian Journal of Agricultural Research 41: 1101–14CrossRefGoogle Scholar
Henderson, A. 1986. A review of pollination studies in the Palmae. Botanical Review 52: 221–59CrossRefGoogle Scholar
Hilje, L. 1984. Fenologia y ecologia floral de Aristolochia grandiflora Swartz (Aristolochiaceae) en Costa Rica. Brenesia 22: 144Google Scholar
Hime, N da C, Costa, E de L. 1985. Sobre Megaselia (M.) aristolochiae n. sp. (Diptera, Phoridae) cujas larvas se criam nas flores de Aristolochia labiata Willd. (Aristolochiaceae). Revista Brasileira Biologia 45: 621–5Google Scholar
Hippa, H., Koponen, S., Osmonen, O. 1981. Pollen transport and pollinating efficiency of flower visitors to the cloudberry (Rubus chamaemorus L.) in northern Fennoscandia. Reports of the Kevo Subarctic Research Station 17: 5866Google Scholar
Hobby, B.M., Smith, K.G.V. 1961. The bionomics of Empis tessellata F. (Diptera: Empididae). Entomologists' Monthly Magazine 97: 210Google Scholar
Hocking, B. 1953. The intrinsic range and flight speed of insects. Transactions of the Royal Entomological Society of London 104: 223345Google Scholar
Hocking, B. 1968. Insect–flower associations in the High Arctic with special reference to nectar. Oikos 19: 359–88CrossRefGoogle Scholar
Hocking, B., Sharplin, C.D. 1965. Flower basking by arctic insects. Nature (London) 206: 215Google Scholar
Hocking, B., Richards, W.R., Twinn, C.R. 1950. Observations on the bionomics of some northern mosquito species (Culicidae: Diptera). Canadian Journal of Research Section D 28: 5880CrossRefGoogle Scholar
Holloway, B.A. 1976. Pollen-feeding in hover-flies (Diptera: Syrphidae). New Zealand Journal of Zoology 3: 339–50CrossRefGoogle Scholar
Horsburgh, M. 1998. A preliminary investigation into the potential pollinators of Symphyotrichum lanceolatum (Willd.) Nesom and Symphyotrichum lateriflorum (L.) Löve & Löve in Waterloo County. M.Sc. dissertation, University of Waterloo, Waterloo, OntarioGoogle Scholar
Hunter, A.S. 1979. New anthophilic Drosophila of Columbia. Annals of the Entomological Society of America 72: 372–83CrossRefGoogle Scholar
Hunter, F.F., Burgin, S.G., Woodhouse, A. 2000. Shattering the folklore: blackflies do not pollinate sweet lowbush blueberry. Canadian Journal of Zoology 78: 2051–4CrossRefGoogle Scholar
Ivri, Y., Dafni, A. 1977. The pollination ecology of Epipactis consimilis Don (Orchidaceae) in Israel. New Phytologist 79: 173–7CrossRefGoogle Scholar
Janzen, D.H. 1979. How to be a fig. Annual Review of Ecology and Systematics 10: 1351CrossRefGoogle Scholar
Johnson, S.D., Steiner, K.E. 1995. Long-proboscid fly pollination of two orchids in the Cape Drakensberg Mountains, South Africa. Plant Systematics and Evolution 195: 169–75CrossRefGoogle Scholar
Johnson, S.D., Steiner, K.E. 1997. Long-tongued fly pollination and evolution of floral spur length in the Disa draconis complex (Orchidaceae). Evolution 51: 4553CrossRefGoogle ScholarPubMed
Karczewski, J. 1967. The observations on flower-visiting species of Tachinidae and Calliphoridae (Diptera). Fragmenta Faunistica (Warsaw) 13: 407 [Polish with English summary]CrossRefGoogle Scholar
Kato, M., Inoue, T., Nagamitsu, T. 1995. Pollination biology of Gnetum (Gnetaceae) in a lowland mixed dipterocarp forest in Sarawak. American Journal of Botany 82: 862–8CrossRefGoogle Scholar
Kearns, C.A. 1992. Anthophilous fly distribution across an elevation gradient. American Midland Naturalist 127: 172–82CrossRefGoogle Scholar
Kearns, C.A., Inouye, D.W. 1993. Techniques for pollination biologists. Niwot, Colo.: University Press of ColoradoGoogle Scholar
Kendall, D.A., Solomon, M.E. 1973. Quantity of pollen on the bodies of insects visiting apple blossom. Journal of Applied Ecology 10: 627–34CrossRefGoogle Scholar
Kevan, P.G. 1970. High arctic insect–flower relations: the inter-relationships of arthropods and flowers at Lake Hazen, Ellesmere Island, Northwest Territories, Canada. PhD dissertation, Department of Entomology, University of Alberta, Edmonton, AlbertaGoogle Scholar
Kevan, P.G. 1972. Insect pollination of High Arctic flowers. Journal of Ecology 60: 831–47CrossRefGoogle Scholar
Kevan, P.G. 1973. Flowers, insects and pollination ecology in the Canadian High Arctic. Polar Record 16: 667–74CrossRefGoogle Scholar
Kevan, P.G. 1989. Thermoregulation in arctic insects and flowers: adaptation and co-adaptation in behaviour, anatomy, and physiology. pp 747–53 in Mercer, J.B. (Ed), Thermal physiology. Amsterdam: Elsevier Science Publishers B.V. (Biomedical Division)Google Scholar
Kevan, P.G. 1990. Sexual differences in temperatures of blossoms on a dioecious plant, Salix arctica: significance for life in the Arctic. Arctic and Alpine Research 22: 283–9CrossRefGoogle Scholar
Kevan, P.G., Baker, H.G. 1983. Insects as flower visitors and pollinators. Annual Review of Entomology 28: 407–53CrossRefGoogle Scholar
Kevan, P.G., Baker, H.G. 1999. Insects on flowers. pp 553–84 in Huffaker, C.B., Gutierrez, A.P. (Eds), Ecological entomology. 2nd ed. New York: John Wiley & SonsGoogle Scholar
Kevan, P.G., Gadawski, R.M., Kevan, S.D., Gadawski, S.E. 1983. Pollination of cranberries, Vaccinium macrocarpon, on cultivated marshes in Ontario. Proceedings of the Entomological Society of Ontario 114: 4553Google Scholar
Kevan, P.G., Tikhmenev, E.A., Usui, M. 1993. Insects and plants in the pollination ecology of the boreal zone. Ecological Research 8: 247–67CrossRefGoogle Scholar
Kingsley, K.J. 1996. Behaviour of the Delhi Sands flower-lving fly (Diptera: Mydidae), a little-known endangered species. Annals of the Entomological Society of America 89: 883–91CrossRefGoogle Scholar
Knab, F. 1907. Mosquitoes as flower visitors. Journal of the New York Entomological Society 15: 215–9Google Scholar
Knight, G.H. 1967. Observations on the behaviour of Bombylius major L., and B. discolor Mik. (Dipt., Bombyliidae) in the midlands. Entomologists' Monthly Magazine 103: 177–81Google Scholar
Knoll, F. 1921. Bombylius fuliginosus und die Farbe der Blumen (Insekten und Blumen I). Abhandlungen Zoologisch-Botanischen Gesellschaft in Wien 12: 17119Google Scholar
Knuth, P. 19061912. Handbook of flower pollination (translated by Davis, JR Ainsworth). Oxford: Oxford University PressGoogle Scholar
Koponen, A. 1990. Entomophily in the Splachnaceae. Botanical Journal of the Linnean Society 104: 115–27CrossRefGoogle Scholar
Krannitz, P.G., Maun, M.A. 1991. Insect visitors to the Guelder Rose, Viburnum opulus var. opulus (Caprifoliaceae), in London, Ont. Canadian Field-Naturalist 105: 13–7CrossRefGoogle Scholar
Kudo, G. 1993. Relationship between flowering time and fruit set of the entomophilous alpine shrub, Rhododendron aureum (Ericaceae), inhabiting snow patches. American Journal of Botany 80: 1300–4CrossRefGoogle Scholar
Kugler, H. 1970. Blutenökologie. Stuttgart: G. FischerGoogle Scholar
Labandeira, C.C. 1997. Insect mouthparts: ascertaining the paleobiology of insect feeding strategies. Annual Review of Ecology and Systematics 28: 153–93CrossRefGoogle Scholar
Labandeira, C.C. 1998 a. How old is the flower and the fly? Science (Washington, DC) 280: 57–9CrossRefGoogle Scholar
Labandeira, C.C. 1998 b. The role of insects in Late Jurassic to middle Cretaceous ecosystems. pp 105–24 in Lucas, S.G., Kirkland, J.I., Estep, J.W. (Eds), Lower and middle Cretaceous terrestrial ecosystems. New Mexico Museum of Natural History and Science Bulletin 14Google Scholar
Lachance, M.A., Starmer, W.T., Rosa, C.A., Bowles, J.M., Barker, J.S.F., Janzen, D.H. 2001. Biogeography of the yeasts of ephemeral flowers and their insects. FEMS Yeast Research 1. In pressGoogle Scholar
Lack, A.J., Diaz, A. 1991. The pollination of Arum maculatum L. — a historical review and new observations. Watsonia 18: 333–42Google Scholar
Lall, S.B., Davies, D.M. 1971. An intergeneric comparison of cephalic structure in tabanids (Diptera) in relation to feeding habits. Journal of Medical Entomology 8: 700–6CrossRefGoogle ScholarPubMed
Leereveld, H. 1982. Anthecological relations between reputedly anemophilous flowers and syrphid flies. III. Worldwide survey of crop and intestine contents of certain anthophilous syrphid flies. Tijdschrift voor Entomologie 125: 2535Google Scholar
Leereveld, H. 1984. Anthecological relations between reputedly anemophilous flowers and syrphid flies. VI. Aspects of the anthecology of Cyperaceae and Sparganium erectum L. Acta Botanica Neerlandica 33: 475–82CrossRefGoogle Scholar
Leereveld, H., Meeuse, A.D.J., Stelleman, P. 1976. Anthecological relations between reputedly anemophilous flowers and syrphid flies. II. Plantago media L. Acta Botanica Neerlandica 25: 205–11CrossRefGoogle Scholar
Leereveld, H., Meeuse, A.D.J., Stelleman, P. 1981. Anthecological relations between reputedly anemophilous flowers and syrphid flies. IV. A note on the anthecology of Scirpus maritimus L. Acta Botanica Neerlandica 30: 465–73CrossRefGoogle Scholar
Leereveld, H., Meeuse, A.D.J., Stelleman, P. 1991. Some cases of visiting of anemophiles by syrphid flies in Madagascar. Israel Journal of Botany 40: 219–23Google Scholar
Levesque, C.M., Burger, J.F. 1982. Insects (Diptera, Hymenoptera) associated with Minuartia groenlandica (Caryophyllaceae) on Mount Washington, New Hampshire, U.S.A., and their possible role as pollinators. Arctic and Alpine Research 14: 117–24CrossRefGoogle Scholar
Lindsey, A.H. 1984. Reproductive biology of Apiaceae. I. Floral visitors to Thaspium and Zizia and their importance in pollination. American Journal of Botany 71: 375–87CrossRefGoogle Scholar
Lorence, D.H. 1985. A monograph of the Monimiaceae (Laurales) in the Malagasy region (southwest Indian Ocean). Annals of the Missouri Botanical Garden 72: 1165CrossRefGoogle Scholar
MacInnes, K.L. 1972. Reproductive ecology of five arctic species of Pedicularis (Scrophulariaceae). Ph.D. dissertation, University of Western Ontario, London, OntarioGoogle Scholar
Magnarelli, L.A. 1978. Nectar-feeding by female mosquitoes and its relation to follicular development and parity. Journal of Medical Entomology 14: 527–30CrossRefGoogle ScholarPubMed
Magnarelli, L.A. 1979. Diurnal nectar-feeding of Aedes cantator and Aedes sollicitans (Diptera: Culicidae). Environmental Entomology 8: 949–55CrossRefGoogle Scholar
Magnarelli, L.A. 1983. Nectar sugars and caloric reserves in natural populations of Aedes canadensis and Aedes stimulans (Diptera: Culicidae). Environmental Entomology 12: 1482–6CrossRefGoogle Scholar
Magnarelli, L.A.. Anderson, J.F., Thorne, J.H. 1979. Diurnal nectar-feeding of salt marsh Tabanidae (Diptera). Environmental Entomology 8: 544–8CrossRefGoogle Scholar
Maier, C.T. 1978. The immature stages and biology of Mallota posticata (Fabricius) (Diptera: Syrphidae). Proceedings of the Entomological Society of Washington 80: 424–40Google Scholar
Manning, J.C., Goldblatt, P. 1996. The Prosoeca peringueyi (Diptera: Nemestrinidae) pollination guild in southern Africa: long-tongued flies and their tubular flowers. Annals of the Missouri Botanical Garden 83: 6786CrossRefGoogle Scholar
Martinez-Ibarra, J.A., Rodriguez, M.H., Arrrendono-Jiminez, J.I., Yuval, B. 1997. Influence of plant abundance on nectar feeding by Aedes aegypti (Diptera: Culicidae) in southern Mexico. Journal of Medical Entomology 34: 589–93CrossRefGoogle ScholarPubMed
McAlpine, J.F. 1965. Observations on anthophilous Diptera at Lake Hazen, Ellesmere Island. Canadian Field-Naturalist 79: 247–52CrossRefGoogle Scholar
McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R., Wood, D.M. 1981. Manual of nearctic Diptera. Vol 1. Agriculture Canada Research Branch Monograph 27Google Scholar
Meeuse, B.J.D., Morris, S. 1984. The sex life of flowers. New York: Facts on File Inc.Google Scholar
Mesler, M.R. 1977. Notes on the floral biology of Plantago in Michigan. Michigan Botanist 16: 7382Google Scholar
Mesler, M.R., Lu, K.L. 1993. Pollination biology of Asarum hartwegii (Aristolochiaceae): an evaluation of Vogel's mushroom-fly hypothesis. Madroño 40: 117–25Google Scholar
Mesler, M.R., Ackerman, J.D., Lu, K.L. 1980. The effectiveness of fungus gnats as pollinators. American Journal of Botany 67: 564–7CrossRefGoogle Scholar
Mosquin, T., Martin, J.E.H. 1967. Observations on the pollination biology of plants on Melville Island, N.W.T., Canada. Canadian Field-Naturalist 81: 201–5CrossRefGoogle Scholar
Motten, A.F. 1986. Pollination ecology of the spring wildflower community of a temperate deciduous forest. Ecological Monographs 56: 2142CrossRefGoogle Scholar
Motten, A.F., Campbell, D.R., Alexander, D.E., Miller, H.L. 1981. Pollination effectiveness of specialist and generalist visitors to a North Carolina population of Claytonia virginica. Ecology 62: 1278–87CrossRefGoogle Scholar
Müller, H. 1883. The fertilisation of flowers. London: Macmillan & Co.CrossRefGoogle Scholar
Nilsson, L.A. 1979. The pollination ecology of Herminium monorchis (Orchidaceae). Botaniska Notiser 132: 537–49Google Scholar
Nilsson, L.A., Jonsson, L., Rason, L., Randrianjohany, E. 1985. Pollination of Plectranthus vestitus (Lamiaceae) by trap-lining hovering bees in Madagascar. Plant Systematics and Evolution 150: 223–36CrossRefGoogle Scholar
Oldroyd, H. 1964. The natural history of flies. London: Weidenfeld and NicolsonGoogle Scholar
Parmenter, L. 1948. Rhingia campestris Mg. (Dipt., Syrphidae) — a further note. Entomologists' Record and Journal of Variation 60: 119–20Google Scholar
Parmenter, L. 1953. The hover flies (Syrphidae). Entomologists' Record and Journal of Variation 65: 122–6, 185–90, 234–8Google Scholar
Parmenter, L. 1956. On Syritta pipiens L. (Syrphidae) and its habits. Entomologists' Record and Journal of Variation 68: 211–4Google Scholar
Parmenter, L. 1958. Flies (Diptera) and their relations with plants. London Naturalist 37: 115–25Google Scholar
Parmenter, L. 1961. Flies visiting the flowers of Wood Spurge, Euphorbia amygdaloides L. [Euphordiaceae (sic)]. Entomologists' Record and Journal of Variation 73: 48–9Google Scholar
Patt, J.M., Merchant, M.W., Williams, D.R.E., Meeuse, B.J.D. 1989. Pollination biology of Platanthera stricta (Orchidaceae) in Olympic National Park, Washington. American Journal of Botany 76: 1097–106CrossRefGoogle Scholar
Patt, J.M., French, J.C., Schal, C., Lech, J., Hartman, T.G. 1995. The pollination biology of tuckahoe, Peltandra virginica (Araceae). American Journal of Botany 82: 1230–40CrossRefGoogle Scholar
Pellmyr, O. 1984. The pollination ecology of Actaea spicata (Ranunculaceae). Nordic Journal of Botany 4: 443–56CrossRefGoogle Scholar
Pellmyr, O. 1989. The cost of mutualism: interactions between Trollius europaeus and its pollinating parasites. Oecologia 78: 53–9CrossRefGoogle ScholarPubMed
Pellmyr, O. 1992. The phylogeny of a mutualism: evolution and coadaptation between Trollius and its seed-parasitic pollinators. Biological Journal of the Linnean Society 47: 337–65CrossRefGoogle Scholar
Philipp, M., Böcher, J., Mattsson, O., Woodell, S.R.J. 1990. A quantitative approach to the sexual reproductive biology and population structure in some arctic flowering plants: Dryas integrifolia, Silene acaulis and Ranunculus nivalis. Meddelelser om Grønland Bioscience 34: 160CrossRefGoogle Scholar
Pont, A.C. 1993. Observations on anthophilous Muscidae and other Diptera (Insecta) in Abisko National Park, Sweden. Journal of Natural History 27: 631–43CrossRefGoogle Scholar
Pratt, G.K., Pratt, H.D. 1972. Records of Tabanidae (Diptera) collected on flowers. Mosquito News 32: 632–3Google Scholar
Primack, R.B. 1978. Variability in New Zealand montane and alpine pollinator assemblages. New Zealand Journal of Ecology 1: 6673Google Scholar
Primack, R.B. 1983. Insect pollination in the New Zealand mountain flora. New Zealand Journal of Botany 21: 317–33CrossRefGoogle Scholar
Prime, C.T. 1960. Lords and ladies. London: CollinsGoogle Scholar
Proctor, M., Yeo, P. 1973. The pollination of flowers. London: CollinsGoogle Scholar
Proctor, M., Yeo, P., Lack, A. 1996. The natural history of pollination. Portland, Oreg.: Timber PressGoogle Scholar
Punchihewa, R.W.K. 1984. Anthophilous insects and the pollination ecology of Asclepias syriaca L., and Asclepias incarnata L. in southern Ontario. M.Sc. dissertation, Department of Environmental Biology, University of Guelph, Guelph, OntarioGoogle Scholar
Raju, A.J.S. 1990. Observations on the floral biology of certain mangroves. Proceedings of the Indian National Science Academy Part B 56: 367–74Google Scholar
Raup, H.M. 1930. The pollination of Habenaria obtusata. Rhodora 32: 88–9Google Scholar
Reader, R.J. 1977. Bog ericad flowers: self-compatibility and relative attractiveness to bees. Canadian Journal of Botany 55: 2279–87CrossRefGoogle Scholar
Reeves, L.M., Reeves, T. 1984. Life history and reproduction of Malaxis paludosa in Minnesota. American Orchid Society Bulletin 53: 1280–91Google Scholar
Ren, D. 1998. Flower-associated Brachycera flies as fossil evidence for Jurassic angiosperm origins. Science (Washington, DC) 280: 85–8CrossRefGoogle Scholar
Richards, A.J. 1986. Plant breeding systems. London: Allen and UnwinGoogle Scholar
Robertson, C. 1924. Flower visits of insects II. Psyche 31: 93111CrossRefGoogle Scholar
Robertson, C. 1928. Flowers and Insects. XXV. Ecology 9: 505–26CrossRefGoogle Scholar
Roubik, D.W. (Editor). 1995. Pollination of cultivated plants in the tropics. Food and Agricultural Organization of the United Nations, Agricultural Services Bulletin 118Google Scholar
Roy, B.A. 1994. The effects of pathogen-induced pseudoflowers and buttercups on each other's insect visitation. Ecology 75: 352–8CrossRefGoogle Scholar
Sabrosky, C.W. 1983. A synopsis of the world species of Desmometopa Loew (Diptera: Milichiidae). Contributions of the Entomological Institute of America 19: 169Google Scholar
Sabrosky, C.W. 1987. A new species of Leptometopa (Diptera, Milichiidae) from Madagascar pollinating Ceropegia (Asclepiadaceae). Proceedings of the Entomological Society of Washington 89: 242–3Google Scholar
Sandholm, H.A., Price, R.D. 1962. Field observations on the nectar feeding habits of some Minnesota mosquitoes. Mosquito News 22: 346–9Google Scholar
Sazima, M. 1978. Polinizacãö por moscas em Bulbophyllum warmingianum Cogn. (Orchidaceae) na Serra do Cipo, Minas Gerais, Brasil. Revista Brasileira de Botanica 1: 133–8Google Scholar
Schemske, D.W., Willson, M.F., Melampy, M.N., Miller, L.J., Verner, L., Schemske, K.M., Best, L.B. 1978. Flowering ecology of some spring woodland herbs. Ecology 59: 351–66CrossRefGoogle Scholar
Schmid, R. 1970. Notes on the reproductive biology of Asterogyne Martiana (Palmae). II. Pollination by syrphid flies. Principes 14: 3949Google Scholar
Schneider, F. 1948. Beitrag zur Kenntnis der Generationsverhältnisse und Diapause räuberischer Schwebfliegen (Syrphidae: Diptera). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 21: 249–88Google Scholar
Schneider, F. 1969. Bionomics and physiology of aphidophagous Syrphidae. Annual Review of Entomology 14: 103–24CrossRefGoogle Scholar
Shaw, D.C., Taylor, R.J. 1986. Pollination ecology of an alpine fell – field community in the North Cascades. Northwest Science 60: 2131Google Scholar
Simes, J.A. 1946. Behaviour of Bombyliidae (Dipt.) while feeding. Entomologists' Monthly Magazine 82: 234Google Scholar
Small, E. 1976. Insect pollinators of the Mer Bleue peat bog of Ottawa. Canadian Field-Naturalist 90: 22–8CrossRefGoogle Scholar
Smart, J. 1943. Simulium feeding on ivy flowers. The Entomologist 76: 2021Google Scholar
Soderstrom, T.R., Calderon, C.E. 1971. Insect pollination in tropical rain forest grasses. Biotropica 3: 116CrossRefGoogle Scholar
Speight, M.C.D. 1978. Flower-visiting flies. pp 229–36 in Stubbs, A., Chandler, P. (Eds), A dipterist's handbook. The Amateur Entomologist, Volume 15. Hanworth: The Amateur Entomologists' SocietyGoogle Scholar
Stebbins, G.L. 1971. California Pipevine: a light trap for unwary flies. California Native Plant Society News-letter 7: 45Google Scholar
Stelleman, P. 1980. Anthecological relations between reputedly anemophilous flowers and syrphid flies. V. Some special aspects of the visiting of Plantago media and P. lanceolata by insects. Beiträge zur Biologie der Pflanzen 55: 157–67Google Scholar
Stelleman, P. 1984. Reflections on the transition from wind pollination to ambophily. Acta Botanica Neerlandica 33: 497508CrossRefGoogle Scholar
Stelleman, P., Meeuse, A.D.J. 1976. Anthecological relations between reputedly anemophilous flowers and syrphid flies. I. The possible role of syrphid flies as pollinators of Plantago. Tijdschrift voor Entomologie 119: 1531Google Scholar
Stoutamire, W.P. 1968. Mosquito pollination of Habenaria obtusata (Orchidaceae). Michigan Botanist 7: 203–12Google Scholar
Stowe, M.K. 1988. Chemical mimicry. pp 513–80 in Spencer, K. (Ed), Chemical mediation of coevolution. London: Academic PressCrossRefGoogle Scholar
Straw, R.M. 1963. Bee-fly pollination of Penstemon ambiguus. Ecology 44: 818–9CrossRefGoogle Scholar
Sugawara, T. 1988. Floral biology of Heterotropa tamaensis (Aristolochiaceae) in Japan. Plant Species Biology 3: 712CrossRefGoogle Scholar
Suvarnayathipat, W. 1984. Mango cultivation. Bangkok: Kasetsart University [in Thai]Google Scholar
Thien, L.B. 1969 a. Mosquito pollination of Habenaria obtusata (Orchidaceae). American Journal of Botany 56: 232–7CrossRefGoogle Scholar
Thien, L.B. 1969 b. Mosquitoes and Habenaria obtusata (Orchidaceae). Mosquito News 29: 252–5Google Scholar
Thien, L.B. 1980. Patterns of pollination in the primitive Angiosperms. Biotropica 12: 113CrossRefGoogle Scholar
Thien, L.B., Utech, F. 1970. The mode of pollination in Habenaria obtusata (Orchidaceae). American Journal of Botany 57: 1031–5CrossRefGoogle Scholar
Thomas, K. 1986. Pollination ecology and reproductive biology of Sarracenia purpurea L. M.Sc. dissertation, York University, Toronto, OntarioGoogle Scholar
Toft, C.A., Kimsey, L.S. 1982. Habitat and behavior of selected Apiocera and Rhaphiomidas (Diptera: Apioceridae), and descriptions of immature stages of A. hispida. Journal of the Kansas Entomological Society 55: 177–86Google Scholar
Totland, O. 1993. Pollination in alpine Norway: flowering phenology, insect visitors, and visitation rates in two plant communities. Canadian Journal of Botany 71: 1072–9CrossRefGoogle Scholar
Troilo, D.B., Cameron, R.G. 1981. Comparative behavior of Pyrellia cyanicolor (Diptera: Muscidae) on the moss Splachuum ampullaceum and on substrates of nutritional value. Great Lakes Entomologist 14: 191–5Google Scholar
Twinn, C.R., Hocking, B., McDuffie, W.C., Cross, H.F. 1948. A preliminary account of the biting flies at Churchill, Manitoba. Canadian Journal of Research Section D 26: 334–57CrossRefGoogle ScholarPubMed
van der Goot, V.S., Grabandt, R.A.J. 1970. Some species of the genera Melanostoma, Platycheirus and Pyrophaena (Diptera: Syrphidae) and their relation to flowers. Entomologische Berichten 30: 135–43Google Scholar
van der Pijl, L. 1953. On the flower biology of some plants from Java, with general remarks on fly traps (species of Annona, Artocarpus, Typhonium, Gnetum, Arisaema, and Abroma). Annales Bogoriensis 1: 7799Google Scholar
van der Pijl, L., Dodson, C.H. 1966. Orchid flowers: their pollination and evolution. Coral Gables, Fla.: University of Miami PressGoogle Scholar
van der Velde, G., Brock, T.C.M., Heine, M., Peeters, PMPM. 1978. Flowers of Dutch Nymphaeaceae as a habitat for insects. Acta Botanica Neerlandica 27: 429–38Google Scholar
Vogel, S. 1954. Blütenbiologische Typen als Elemente der Sippengliederung: dargestellt anhand der Flora Süd Afrikas. Botanical Studien 1: 1338Google Scholar
Vogel, S. 1978. Pilzmuckenblumen als Pilzmimeten. [Fungus-gnat flowers mimicking fungi.] Flora 167: 329–98CrossRefGoogle Scholar
Vogt, C.A. 1990. Pollination in Cypripedium reginae (Orchidaceae). Lindleyana 5: 145–50Google Scholar
Voss, E.G., Riefner, R.E. 1983. A pyralid moth (Lepidoptera) as pollinator of blunt-leaf orchid. Great Lakes Entomologist 16: 5760Google Scholar
Waitzbauer, V.W. 1976. Die Insektenfauna männlicher Blutenstande von Typha angustifolia. Zoologischer Anzeiger 196: 915Google Scholar
Warncke, E., Terndrup, U., Michelsen, V., Erhardt, A. 1993. Flower visitors to Saxifraga hirculus in Switzerland and Denmark, a comparative study. Botanica Helvetica 103: 141–7Google Scholar
Weemns, H.V. Jr. 1953. Notes on collecting syrphid flies (Diptera: Syrphidae). Florida Entomologist 36: 91–8CrossRefGoogle Scholar
Whitehead, V.B., Giliomee, J.H., Rebelo, A.G. 1987. Insect pollination in the Cape flora. pp 5282in Rebelo, A.G. (Ed), A preliminary synthesis of pollination biology in the Cape flora. South African National Science Programmes Report 141Google Scholar
Willemstein, S.C. 1987. An evolutionary basis for pollination ecology. Leiden: E.J. Brill/Leiden University PressCrossRefGoogle Scholar
Williams, A.H. 1995. Adult female Mydas clavatus (Diptera: Mydidae) feeding on flowers in Wisconsin. Great Lakes Entomologist 28: 227–9Google Scholar
Willis, J.C., Burkill, I.H. 18951908. Flowers and insects in Great Britain. I–IV. Annals of Botany 9: 227–73; 17: 313–49, 539–70; 22: 603–49CrossRefGoogle Scholar
Wilson, B.H., Lieux, M. 1972. Pollen grains in the guts of field collected tabanids in Louisiana. Annals of the Entomological Society of America 65: 1264–6CrossRefGoogle Scholar
Wirth, W.W. 1991. New and little-known species of Forcipomyia (Diptera: Ceratopogonidae) associated with cocoa pollination in Brazil. Proceedings of the Entomological Society of Washington 93: 163–75Google Scholar
Wolda, H., Sabrosky, C.W. 1986. Insect visitors to two forms of Aristolochia pilosa in Las Cumbres, Panama. Biotropica 18: 295–9CrossRefGoogle Scholar
Yafuso, M. 1993. Thermogenesis of Alocasia odora (Araceae) and the role of Colocasiomyia flies (Diptera: Drosophilidae) as cross-pollinators. Environmental Entomology 22: 601–6CrossRefGoogle Scholar
Young, A.M. 1984 a. Ecological notes on cacao-associated midges (Diptera: Ceratopogonidae) in the “Catongo” cacao plantation at Turrialba, Costa Rica. Proceedings of the Entomological Society of Washington 86: 185–94Google Scholar
Young, A.M. 1984 b. Mechanism of pollination by Phoridae (Diptera) in some Herrania species (Sterculiaceae) in Costa Rica. Proceedings of the Entomological Society of Washington 86: 503–18Google Scholar
Young, A.M. 1985. Studies of cecidomyiid midges (Diptera: Cecidomyiidae) as cocoa pollinators (Theobroma cacao) in Central America. Proceedings of the Entomological Society of Washington 87: 4979Google Scholar
Young, A.M. 1986. Cocoa pollination. Cocoa Grower's Bulletin 37: 523Google Scholar