Skip to main content
SpringerLink
Log in

Linearly Polarized Light as a Guiding Cue for Water Detection and Host Finding in Tabanid Flies

  • Chapter
  • First Online:
Polarized Light and Polarization Vision in Animal Sciences

Part of the book series: Springer Series in Vision Research ((SSVR,volume 2))

Abstract

In this chapter we show that tabanid flies are attracted to horizontally polarized light stimulating their ventral eye region. Female and male tabanids use this polarotaxis governed by the horizontal E-vector to find water, while another type of polarotaxis based on the degree of polarization serves host finding by female tabanids. We show that female tabanids are less attracted to bright than dark hosts, the reason for which is partly that dark hosts reflect light with higher degrees of polarization than bright hosts. We also demonstrate that the use of a striped fur pattern has the advantage that such coat patterns attract far fewer tabanids than either homogeneous black, brown, grey or white equivalents. The attractiveness of striped patterns to tabanids is also reduced if only polarization modulations (parallel stripes with alternating orthogonal directions of polarization) occur in homogeneous grey surfaces. The attractiveness to tabanids decreases with decreasing stripe width, and stripes below a certain width threshold are unattractive at all to tabanids. Further, the stripe widths of zebra coats fall in a range where the striped pattern is most unattractive to tabanids. Tabanids are strongly attracted by CO2 and ammonia emitted by their hosts. We show here that the poor visual attractivity of stripes to tabanids is not overcome by olfactory attractiveness. Finally, we show that dark spots on a bright coat surface also disrupt the visual attractiveness to tabanids. The smaller and the more numerous the spots, the less attractive the host is to tabanids. The attractiveness of spotty patterns to tabanids is also reduced if the target exhibits spottiness only in the angle of polarization pattern, while being homogeneous grey with a constant high degree of polarization. This could be one of the possible evolutionary benefits that explains why spotty coat patterns are so widespread in mammals, especially in ungulates, many species of which are tabanid hosts.

This is a preview of subscription content, log in via an institution to check access.

Access this chapter

Log in via an institution
Chapter
USD 29.95
Price excludes VAT (USA)
  • Available as PDF
  • Read on any device
  • Instant download
  • Own it forever
eBook
USD 129.00
Price excludes VAT (USA)
  • Available as EPUB and PDF
  • Read on any device
  • Instant download
  • Own it forever
Softcover Book
USD 169.99
Price excludes VAT (USA)
  • Compact, lightweight edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info
Hardcover Book
USD 169.99
Price excludes VAT (USA)
  • Durable hardcover edition
  • Dispatched in 3 to 5 business days
  • Free shipping worldwide - see info

Tax calculation will be finalised at checkout

Purchases are for personal use only

Institutional subscriptions

Notes

  1. 1.

    It is well known among horse keepers that in shady refuges, in forests or in stables, horses suffer tabanid annoyance only rarely. Tabanid flies do not follow horses and other host animals in large numbers to such shady places more or less surrounded by vegetation or artificial walls. The reasons for this are the following (Horváth et al. 2010b): (1) Tabanids need enough free, open space to fly. (2) Many tabanid species need a higher body temperature to start flying so quickly that they can successfully escape from the defensive reactions of a host animal (e.g. tail swishing, biting, licking), by which the host tries to drive them away from its body. In shady refuges the wing muscles of tabanids may not function rapidly enough for such an escape. Therefore, tabanids attack host animals usually under sunny conditions. For the same reason, tabanids usually do not fly and attack hosts on overcast days and under rainy or too windy conditions. However, certain small-sized tabanids prefer periods prior to rain, because the lower air temperature and the higher air humidity are advantageous to these insects, whose body has a large surface/volume ratio, and thus they can easily become dehydrated under sunny, dry and windy conditions. (3) The forest vegetation, as a structured background, makes more difficult the visual recognition of host animals.

    On the other hand, when looking for water by means of the horizontally polarized water-reflected light, tabanids can quickly fly through shady areas and touch the water surface to drink and/or bath, for example. This water-seeking behaviour of male and female tabanids is quite different from the blood-sucking behaviour of tabanid females. The former can happen under both shady and sunny conditions, while the latter is usually performed in sunshine.

References

  • Allan SA, Day JF, Edman JD (1987) Visual ecology of biting flies. Annu Rev Entomol 32:297–316

    Article  PubMed  CAS  Google Scholar 

  • Allen WL, Cuthill IC, Scott-Samuel NE, Baddeley R (2010) Why the leopard got its spots: relating pattern development to ecology in felids. Proc R Soc B 278:1373–1380

    Article  PubMed  PubMed Central  Google Scholar 

  • Baldacchino F, Desquesnes M, Mihok S, Foil LD, Duvallet G, Jittapalapong S (2014) Tabanids: Neglected subjects of research, but important vectors of disease agents! Infection, Genetics and Evolution (in press). doi:10.1016/j.meegid.2014.03.029

  • Bard JBL (1977) A unity underlying the different zebra striping patterns. J Zool (Lond) 183:527–539

    Google Scholar 

  • Becker CD, Ginsberg JR (1990) Mother-infant behaviour of wild Grevy’s zebra: adaptations for survival in semi-desert East Africa. Anim Behav 40:1111–1118

    Article  Google Scholar 

  • Blahó M, Egri Á, Báhidszki L, Kriska G, Hegedüs R, Åkesson S, Horváth G (2012) Spottier targets are less attractive to tabanid flies: on the tabanid-repellency of spotty fur patterns. PLoS One 7(8):e41138. doi:10.1371/journal.pone.0041138 + supporting information

  • Blahó M, Egri Á, Száz D, Kriska G, Åkesson S, Horváth G (2013) Stripes disrupt odour attractiveness to biting horseflies: battle between ammonia, CO2, and colour pattern for dominance in the sensory systems of host-seeking tabanids. Physiol Behav 119:168–174

    Article  PubMed  Google Scholar 

  • Brady J, Shereni A (1988) Landing responses of the tsetse fly Glossina norsitans norsitans Weidemann and the stable fly Stomoxys calcitrans (L.) (Diptera: Glossinidae and Muscidae) to black and white patterns: a laboratory study. Bull Entomol Res 78:301–311

    Article  Google Scholar 

  • Brega J (2005) Essential equine studies: anatomy and physiology. JA Allen and Co Ltd, London

    Google Scholar 

  • Burakova O, Mazokhin-Porshnyakov G (1982) Electron microscopy of the compound eye in Haematopota pluvialis L. (Diptera: Tabanidae). Ent Rev 61:26–33

    Google Scholar 

  • Caro T (2009) Contrasting coloration in terrestrial mammals. Philos Trans R Soc B 364:537–548

    Article  Google Scholar 

  • Caro T, Izzo A, Reiner RC, Walker H, Stankowich T (2014) The function of zebra stripes. Nature Communications 5:3535. doi:10.1038/ncomms4535

  • Cloudsley-Thompson JL (1984) How the zebra got his stripes—new solutions to an old problem. Biologist 31:226–228

    Google Scholar 

  • Cott HB (1957) Adaptive colouration in animals. John Dickens, Northampton

    Google Scholar 

  • Cott HB (1966) Colouration in animals. Methuen, London

    Google Scholar 

  • Csabai Z, Boda P, Bernáth B, Kriska G, Horváth G (2006) A ‘polarisation sun-dial’ dictates the optimal time of day for dispersal by flying aquatic insects. Freshw Biol 51:1341–1350

    Article  Google Scholar 

  • Darwin CR (1871) The descent of man, and selection in relation to sex, vol 2. John Murray, London, p 302

    Book  Google Scholar 

  • Duncan P, Vigne N (1979) The effect of group size in horses on the rate of attacks by blood-sucking flies. Anim Behav 27:623–625

    Article  Google Scholar 

  • Egri Á, Blahó M, Kriska G, Farkas R, Gyurkovszky M, Åkesson S, Horváth G (2012a) Polarotactic tabanids find striped patterns with brightness and/or polarization modulation least attractive: an advantage of zebra stripes. J Exp Biol 215: 736–745 + electronic supplement

    Google Scholar 

  • Egri Á, Blahó M, Sándor A, Kriska G, Gyurkovszky M, Farkas R, Horváth G (2012b) New kind of polarotaxis governed by degree of polarization: Attraction of tabanid flies to differently polarizing host animals and water surfaces. Naturwissenschaften 99:407–416 + electronic supplement

    Google Scholar 

  • Eltringham SK (1979) The ecology and conservation of large African mammals. Macmillan, Basingstoke

    Google Scholar 

  • Estes RD (1992) The behavior guide to african mammals including hoofed mammals, carnivores, primates. University of California Press, Los Angeles, CA

    Google Scholar 

  • Finch VA, Bennett IL, Holmes CR (1984) Coat colour in cattle: effect on thermal balance, behaviour and growth, and relationship with coat type. J Agric Sci 102:141–147

    Article  Google Scholar 

  • Foil LD (1989) Tabanids as vectors of disease agents. Parasitol Today 5:88–96

    Article  PubMed  CAS  Google Scholar 

  • Galton F (1851) South Africa. Minerva Library, New York

    Google Scholar 

  • Gibson G (1992) Do tsetse-flies ‘see’ zebras? A field study of the visual response of tsetse to striped targets. Physiol Entomol 17:141–147

    Article  Google Scholar 

  • Glenn EW (1983) Coat color and solar heat gain in animals. Bioscience 33:88–91

    Article  Google Scholar 

  • Godfrey D, Lythgoe JN, Rumball DA (1987) Zebra stripes and tiger stripes: the spatial frequency distribution of the pattern compared to that of the background is significant in display and crypsis. Biol J Linn Soc 32:427–433

    Article  Google Scholar 

  • Hall MJR, Farkas R, Chainey JE (1998) Use of odour-baited sticky boards to trap tabanid flies and investigate repellents. Med Vet Entomol 12:241–245

    Article  PubMed  CAS  Google Scholar 

  • Harris RHTP (1930) Report on the bionomics of the tsetse fly. Provincial Administration of Natal, Pietermaritzburg, South Africa

    Google Scholar 

  • Hayakawa H (1980) Biological studies on Tabanus iyoensis group of Japan, with special reference to their blood-sucking habits (Diptera, Tabanidae). Bull Tohoku Nat Agric Exp Station 62:131–321

    Google Scholar 

  • Horváth G, Kriska G (2008) Polarization vision in aquatic insects and ecological traps for polarotactic insects. In: Lancaster J, Briers RA (eds) Aquatic insects: challenges to populations. CAB International Publishing, Wallingford, Oxon, pp 204–229, Chapter 11

    Google Scholar 

  • Horváth G, Varjú D (2004) Polarized light in animal vision—polarization patterns in nature. Springer, Heidelberg

    Book  Google Scholar 

  • Horváth G, Malik P, Kriska G, Wildermuth H (2007) Ecological traps for dragonflies in a cemetery: the attraction of Sympetrum species (Odonata: Libellulidae) by horizontally polarizing black gravestones. Freshw Biol 52:1700–1709

    Article  Google Scholar 

  • Horváth G, Majer J, Horváth L, Szivák I, Kriska G (2008) Ventral polarization vision in tabanids: horseflies and deerflies (Diptera: Tabanidae) are attracted to horizontally polarized light. Naturwissenschaften 95:1093–1100

    Article  PubMed  Google Scholar 

  • Horváth G, Blahó M, Egri Á, Kriska G, Seres I, Robertson B (2010a) Reducing the maladaptive attractiveness of solar panels to polarotactic insects. Conserv Biol 24:1644–1653 + electronic supplement

    Google Scholar 

  • Horváth G, Blahó M, Kriska G, Hegedüs R, Gerics B, Farkas R, Åkesson S (2010b) An unexpected advantage of whiteness in horses: the most horsefly-proof horse has a depolarizing white coat. Proc R Soc B 277:1643–1650

    Article  PubMed  PubMed Central  Google Scholar 

  • Horváth G, Móra A, Bernáth B, Kriska G (2011) Polarotaxis in non-biting midges: female chironomids are attracted to horizontally polarized light. Physiol Behav 104:1010–1015 + cover picture

    Google Scholar 

  • Hribar LJ, LePrince DJ, Foil LD (1992) Ammonia as an attractant for adult Hybomitra lasiophthalma (Diptera: Tabanidae). J Med Entomol 29:346–348

    PubMed  CAS  Google Scholar 

  • Jékely G (2009) Evolution of phototaxis. Philos Trans R Soc B 364:2795–2808

    Article  Google Scholar 

  • Jordan AM (1986) Trypanosomiasis control and African rural development. Longman, New York

    Google Scholar 

  • Kingdon J (1984) The zebra’s stripes: an aid to group cohesion. In: MacDonald D (ed) The encyclopaedia of mammals. Equinox, Oxford, pp 486–487

    Google Scholar 

  • Kipling R (1908) Just so stories. Macmillan, London

    Google Scholar 

  • Krcmar S (2005a) Seasonal abundance of horse flies (Diptera: Tabanidae) from two locations in eastern Croatia. Journal of Vector Ecology 30:316–321

    Google Scholar 

  • Krcmar S (2005b) Response of horse flies (Diptera, Tabanidae) to different olfactory attractants. Biologia Bratislava 60:611–613

    Google Scholar 

  • Krcmar S (2013) Comparison of the efficiency of the olfactory and visual traps in the collection of horseflies (Diptera: Tabanidae). Entomologia Generalis 34:261–267

    Google Scholar 

  • Krcmar S, Maric S (2006) Analysis of the feeding sites for some horse flies (Diptera, Tabanidae) on a human in Croatia. Collegium Antropologicum 30:901–904

    Google Scholar 

  • Krcmar S, Merdic E, Kopi M (2005) Diurnal periodicity in the biting activity of horsefly species in the Kopacki rit Nature Park, Croatia (Diptera: Tabanidae). Entomologia Generalis 28:139–146

    Google Scholar 

  • Kriska G, Csabai Z, Boda P, Malik P, Horváth G (2006) Why do red and dark-coloured cars lure aquatic insects? The attraction of water insects to car paintwork explained by reflection-polarization signals. Proc R Soc B 273:1667–1671

    Article  PubMed  PubMed Central  Google Scholar 

  • Kriska G, Bernáth B, Horváth G (2007) Positive polarotaxis in a mayfly that never leaves the water surface: polarotactic water detection in Palingenia longicauda (Ephemeroptera). Naturwissenschaften 94:148–154 + cover picture

    Google Scholar 

  • Kriska G, Malik P, Szivák I, Horváth G (2008) Glass buildings on river banks as “polarized light traps” for mass-swarming polarotactic caddis flies. Naturwissenschaften 95:461–467

    Article  PubMed  CAS  Google Scholar 

  • Kriska G, Bernáth B, Farkas R, Horváth G (2009) Degrees of polarization of reflected light eliciting polarotaxis in dragonflies (Odonata), mayflies (Ephemeroptera) and tabanid flies (Tabanidae). J Insect Physiol 55:1167–1173

    Article  PubMed  CAS  Google Scholar 

  • Kruuk H (1972) The spotted hyena. University of Chicago Press, Chicago, IL

    Google Scholar 

  • Lehane MJ (2005) The biology of blood-sucking in insects, 2nd edn. Cambridge University Press, Cambridge, UK

    Book  Google Scholar 

  • Lerner A, Meltser N, Sapir N, Erlick C, Shashar N, Broza M (2008) Reflected polarization guides chironomid females to oviposition sites. J Exp Biol 211:3536–3543

    Article  PubMed  Google Scholar 

  • Louw GN (1993) Physiological animal ecology. Longman, New York

    Google Scholar 

  • Majer J (1987) Tabanids—Tabanidae. In: Szekessy V (ed) Fauna Hungariae, vol 14(9). Akadémiai Kiadó, Budapest, pp 1–57 (in Hungarian)

    Google Scholar 

  • Marler P, Hamilton WJ (1968) Mechanisms of animal behavior. Wiley, New York

    Google Scholar 

  • Marlin D, Nankervis KJ (2002) Equine exercise physiology. Wiley-Blackwell, New York

    Google Scholar 

  • McLeod DNK (1987) Zebra stripes. New Sci 115:68

    Google Scholar 

  • Menzel R (1979) Spectral sensitivity and colour vision in invertebrates. In: Autrum H (ed) Comparative physiology and evolution of vision in invertebrates, vol VII/6A. Springer, Berlin, pp 503–580

    Chapter  Google Scholar 

  • Mihok S (2002) The development of a multipurpose trap (the Nzi) for tsetse and other biting flies. Bull Entomol Res 92:385–403

    Article  PubMed  CAS  Google Scholar 

  • Mihok S, Lange K (2012) Synergism between ammonia and phenols for Hybomitra tabanids in northern and temperate Canada. Med Vet Entomol 26:282–290

    Article  PubMed  CAS  Google Scholar 

  • Mihok S, Mulye H (2010) Responses of tabanids to Nzi traps baited with octenol, cow urine and phenols in Canada. Med Vet Entomol 24:266–272

    PubMed  CAS  Google Scholar 

  • Morris D (1990) Animal watching. A field guide to animal behaviour. Jonathan Crape, London

    Google Scholar 

  • Ortolani A, Caro TM (1996) The adaptive significance of color patterns in carnivores: phylogenetic tests of classic hypotheses. In: Gittleman J (ed) Carnivore behaviour, ecology and evolution, vol 2, Cornell University Press. Ithaca, NY, pp 132–188

    Google Scholar 

  • Pielberg GR, Golovko A, Sundström E, Curik I, Lennartsson J, Seltenhammer MH, Druml T, Binns M, Fitzsimmons C, Lindgren G, Sandberg K, Baumung R, Vetterlein M, Strömberg S, Grabherr M, Wade C, Lindblad-Toh K, Pontén F, Heldin CH, Sölkner J, Andersson L (2008) A cis-acting regulatory mutation causes premature hair graying and susceptibility to melanoma in the horse. Nat Genet 40:1004–1009

    Article  Google Scholar 

  • Prothero DR, Schoch RM (2003) Horns, tusks, and flippers: the evolution of hoofed mammals. Johns Hopkins University Press, Baltimore, MD

    Google Scholar 

  • Rutberg AT (1987) Horse fly harassment and the social behaviour of feral ponies. Ethology 75:145–154

    Article  Google Scholar 

  • Ruxton GD (2002) The possible fitness benefits of striped coat coloration for zebra. Mammal Rev 32:237–244

    Article  Google Scholar 

  • Schwind R (1983) A polarization-sensitive response of the flying water bug Notonecta glauca to UV light. J Comp Physiol 150:87–91

    Article  Google Scholar 

  • Schwind R (1984) Evidence for true polarization vision based on a two-channel analyser system in the eye of the water bug, Notonecta glauca. J Comp Physiol A 154:53–57

    Article  Google Scholar 

  • Schwind R (1985a) A further proof of polarization vision of Notonecta glauca and a note on threshold intensity for eliciting the plunge reaction. Experientia 41:466–467

    Article  Google Scholar 

  • Schwind R (1985b) Sehen unter und über Wasser, Sehen von Wasser. Naturwissenschaften 72:343–352

    Article  Google Scholar 

  • Schwind R (1989) A variety of insects are attracted to water by reflected polarized light. Naturwissenschaften 76:377–378

    Article  Google Scholar 

  • Schwind R (1991) Polarization vision in water insects and insects living on a moist substrate. J Comp Physiol A 169:531–540

    Article  Google Scholar 

  • Schwind R (1995) Spectral regions in which aquatic insects see reflected polarized light. J Comp Physiol A 177:439–448

    Article  Google Scholar 

  • Schwind R (1999) Daphnia pulex swims towards the most strongly polarized light—a response that leads to ‘shore flight’. J Exp Biol 202:3631–3635

    PubMed  Google Scholar 

  • Smith WC, Butler JF (1991) Ultrastructure of the Tabanidae compound eye: unusual features for Diptera. J Insect Physiol 37:287–296

    Article  Google Scholar 

  • Stevens M, Merilaita S (2009) Animal camouflage: current issues and new perspectives. Philos Trans R Soc B 364:423–427

    Article  Google Scholar 

  • Tashiro H, Schwardt HH (1953) Biological studies of horseflies in New York. J Econ Entomology 46:813–822

    Google Scholar 

  • Thayer AH (1909) Concealing coloration in the animal kingdom. Macmillan, New York

    Google Scholar 

  • Thorsteinson AJ, Bracken GK, Hanec W (1965) The orientation behaviour of horseflies and deerflies (Tabanidae: Diptera). III. The use of traps in the study of orientation of tabanids in the field. Entomol Exp Appl 8:189–192

    Article  Google Scholar 

  • Tresidder J (ed) (2005) The complete dictionary of symbols. Chronicle Books, San Francisco, CA

    Google Scholar 

  • Turner DA, Invest JF (1973) Laboratory analyses of vision in tsetse flies (Dipt. Glossinidae). Bull Entomol Res 62:343–357

    Article  Google Scholar 

  • Vale GA (1974) The response of tsetse flies (Diptera, Glossinidae) to mobile and stationary baits. Bull Entomol Res 64:545–588

    Article  Google Scholar 

  • Vaughan TA (1986) Mammology, 3rd edn. North Arizona University, Flagstaff, AZ

    Google Scholar 

  • Waage JK (1981) How the zebra got its stripes—biting flies as selective agents in the evolution of zebra coloration. J Entomol Soc South Afr 44:351–358

    Google Scholar 

  • Wallace AR (1867) Mimicry and other protective resemblances among animals. Westm Foreign Q Rev 32:1–43

    Google Scholar 

  • Wallace AR (1879) The protective colours of animals. Science 2:128–137

    Google Scholar 

  • Wildermuth H (1998) Dragonflies recognize the water of rendezvous and oviposition sites by horizontally polarized light: a behavioural field test. Naturwissenschaften 85:297–302

    Article  CAS  Google Scholar 

  • Wilson BH, Tugwell NP, Burns EC (1966) Attraction of tabanids to traps baited with dry-ice under field conditions in Louisiana. J Med Entomol 3:148–149

    Google Scholar 

Download references

Author information

Authors and Affiliations

  1. Environmental Optics Laboratory, Department of Biological Physics, Physical Institute, Eötvös University, Pázmány sétány 1, 1117, Budapest, Hungary

    Gábor Horváth, Ádám Egri & Miklós Blahó

Authors
  1. Gábor Horváth
    View author publications

    You can also search for this author in PubMed Google Scholar

  2. Ádám Egri
    View author publications

    You can also search for this author in PubMed Google Scholar

  3. Miklós Blahó
    View author publications

    You can also search for this author in PubMed Google Scholar

Corresponding author

Correspondence to Gábor Horváth .

Editor information

Editors and Affiliations

  1. Environmental Optics Laboratory, Dept. Biological Physics, Eötvös University, Budapest, Hungary

    Gábor Horváth

1 Electronic Supplementary Material

Supplementary figures and one video clip are available in the online version of this chapter. The videos can also be accessed at http://www.springerimages.com/videos/ <ISBN print>.

71484_2_En_22_MOESM60_ESM.zip

Typical water-touching behaviour of a polarotactic tabanid fly at a horizontal shiny black test surface in a cemetery in Kiskunhalas (Hungary) on 11 July 2006. The touchdowns of the tabanid were elicited by the strongly and horizontally polarized light reflected from the test surface (copyright holders: Dr. Gábor Horváth and Loránd Horváth) (AVI 25123 kb)

Rights and permissions

Reprints and permissions

Copyright information

© 2014 Springer-Verlag Berlin Heidelberg

About this chapter

Cite this chapter

Horváth, G., Egri, Á., Blahó, M. (2014). Linearly Polarized Light as a Guiding Cue for Water Detection and Host Finding in Tabanid Flies. In: Horváth, G. (eds) Polarized Light and Polarization Vision in Animal Sciences. Springer Series in Vision Research, vol 2. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-642-54718-8_22

Download citation

Publish with us

Policies and ethics

Search

Navigation