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Why
are traps blue - When there are no blue animals?
It is hard to
understand why
biting flies investigate
simple, artificial objects in the
absence of appropriate host odours. However, from the
kinds of flies captured,
and their blood-feeding habits
(females only in many species),
most biting flies likely perceive traps as an
opportunity to obtain a blood meal.
Some species also investigate objects to encounter the opposite sex
(males of many species), to rest in the sun or shade depending on the
temperature (perhaps all species?), or to simply get out of the wind (stable flies).
As a
general rule, flies react best
to artificial objects
(traps or targets) that are about the size of
large animals, and that
contrast well with
their surroundings. Some species are
very sensitive to
movement, and hardly react to stationary objects (deer flies (Chrysops),
and savannah tsetse such as G. morsitans, G. swynnertoni). A
universal explanation for why blue and black traps catch large numbers of
biting flies is unrealistic, given the variety of patterns observed.
A recent
publication has suggested that tsetse in particular are attracted to blue
objects simply because
of the tendency of daytime shadows to be both dark and blue at the same
time. This is a reasonable hypothesis for many observations, and is worth testing with appropriate experimental
studies.
Steverding, D. & Troscianki, T. (2004) On the role of
blue shadows in the visual behaviour of tsetse flies. Proceedings of the
Royal Society, Biological Sciences, Biology Letters 271, S16-S17.
PDF file |
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Biting flies perceive the environment differently
than
people. Their eyes respond to
ultraviolet wavelengths, and they
have a greatly reduced ability to discern fine detail.
Our retinas have peak
sensitivity at about 550 nm
(yellow). In
our visible range (400-700
nm), flies see "colours" at low wavelengths (violet and blue),
with some sensitivity in the
blue-green. Biting flies cannot see red, although some
other insects can.
Biting flies perceive the ultraviolet
exceptionally well (<400 nm).
Hence, unlike our primary "trichromatic" visual response to
blue, green and red, flies likely see primary "colours" corresponding to ultraviolet,
blue and blue-green. A
highly-visible object for a
biting fly might therefore be
an object that contrasts
well at low wavelengths relative to green vegetation.
There is considerable literature on the perception of colour in certain
insects; the two papers below are a good introduction.
Briscoe, A.D. & Chittka, L. (2001) The evolution of
color vision in insects. Annual Review of Entomology 46, 471-510.
Stavenga, D. (2002) Colour in the eyes of insects. Journal of
Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral
Physiology, 188, 337-348.
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A simple explanation for "why traps are
blue" is illustrated below. The graph shows the ratio of
reflected light for two arbitrary kinds of blue cloth relative to the kinds of vegetation
found where tsetse live in Africa (bright green Acacia
leaves on a dark green Combretum leaf). Note how a dark blue cotton
still appears as a bright object in the blue region at 450 nm relative to
vegetation (reflecting 300% as
much light). When combined with black cloth, a blue trap would be a novel object,
perhaps worthy of investigation for a hungry fly. To us,
a bright blue polyester would look similar to the
blue cotton, but to a fly, it would look very different. We
would notice the additional brightness in the blue
region, but the fly would be able to detect the large difference in
reflectance in the ultraviolet - violet. Both pieces of cloth would appear as
bright objects to the fly, but the fly would see a more complex "colour" in the polyester. The smooth
surface of the polyester would also cause polarisation of
reflected light. The fly would also be able to see this
feature. |
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These
standard reflectance data are for an
artificial situation in the laboratory (new, dry
cloth against a black background). Economical trap
fabrics are often not opaque, and hence they will both reflect and transmit
light under natural conditions. Cloth
also appears different
when wet or weathered, when in the sun, cloud or shade, when viewed
against growing or senescent vegetation, etc. Changes in
appearance also occur as the sun's angle changes
during the day,
resulting in differing amounts of light falling
directly on the trap, or
reflected onto it from the
environment. These "real-world" features
make it difficult to predict how any fabric will perform in nature, based
solely on standard reflectance data obtained in the laboratory.
To illustrate these
phenomena, I have taken a some representative photographs of traps in
nature, using blue and ultraviolet filters. Follow the link below to see
how the environment might look through the eyes of a biting fly.
Traps in
Nature
(MANY IMAGES, PAGE LOADS SLOWLY)
For a technical introduction to
the many factors involved in the description and measurement of colour
see:
Nassau, K. (1998) Color for science, art and technology.
Amsterdam, Elsevier. |
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Biting Fly Spectral Sensitivity
Below are some examples of spectral
sensitivity measurements (electroretinograms) for biting flies. Note that
these graphs are on different scales and are based on different
experimental criteria. Also note that sensitivity does not simply
translate into a readily-observable behaviour. There are many differences
among species in terms of how they are attracted by, orient towards, and
land on, objects of different colours (Visual
Bibliography). |
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Green, C.H. &
Cosens, D. (1983) Spectral responses of the tsetse fly Glossina
morsitans morsitans. Journal of Insect Physiology 29,
795-800. |
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Agee, H.R. &
Patterson, R.S. (1983) Spectral sensitivity of stable, face, and horn
flies and behavioral responses of stable flies to visual traps (Diptera
Muscidae). Environmental Entomology 12, 1823-1827. |
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Allan, S.A.,
Stoffolano, J.G., Jr. & Bennett, R.R. (1991) Spectral sensitivity of
the horse fly Tabanus nigrovittatus (Diptera Tabanidae).
Canadian Journal of Zoology 69, 369-374. |
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