Approximately 1 day after eclosion, the imago of the
hummingbird hawkmoth Macroglossum stellatarum starts
searching for nectar sources. How does it find the first flower?
Like other flower-visiting insects, it must be guided by means
of an innate flower template (Darwin, 1877). The template may
contain various features, such as odour (see Knoll, 1925, 1926;
Ilse, 1928), colour (Lepidoptera: Ilse, 1928; Swihart, 1971;
Scherer and Kolb, 1987a,b; Cutler et al. 1995; bumblebees:
Lunau, 1990; Lunau et al. 1996; honeybees: Giurfa et al.
1995b; hoverflies: Ilse, 1949; Lunau and Wacht, 1994) and
flower size and pattern (Lepidoptera: Ilse, 1933; honeybees:
Lehrer et al. 1995).
Analysing spontaneous preferences is interesting for two
main reasons. First, it is a very useful method for studying
animals that cannot easily be trained to a particular stimulus,
such as crickets (Campan and Lacoste, 1971) and beetles
(Varjú, 1976; Dafni, 1997). In several lepidopterans that
cannot easily be trained to flower colours, the feeding response
has been described using this technique as a wavelength-
specific behaviour, the sensitivity of which depends mostly on
the blue, and to a lesser extent on the green, receptor (Scherer
and Kolb, 1987a,b; Cutler et al. 1995). In different contexts,
the same animals might perform different wavelength-specific
behaviours or even demonstrate true colour vision, as has been
shown in the case of the honeybee (Menzel, 1979). Second,
spontaneous choices might tell us which sensory cues are
behaviourally important in the environment. Giurfa et al.
(1995b) gave an ecological explanation for innate colour
preferences: they have shown that honeybees are guided to
flowers that promise a large amount of nectar. Lehrer et al.
(1995) analysed pattern preferences in the same insect and
concluded that flower-like patterns are preferred to others.
They did not, however, work with flower-naive bees.
The day-active hummingbird hawkmoth Macroglossum
stellatarum is attracted by visual stimuli alone, although
odours may also influence its motivational state (Brantjes,
1973). In contrast to other species (Knoll, 1926; Ilse, 1928;
Brantjes, 1973; Scherer and Kolb, 1987b), this influence is not
necessary to induce feeding behaviour in M. stellatarum
(Knoll, 1922). Despite the fact that no electrophysiological
recordings from its photoreceptors exist, it can be assumed that
M. stellatarum has three spectral receptor types with
827 The Journal of Experimental Biology 200, 827–836 (1997)
Printed in Great Britain © The Company of Biologists Limited 1997
JEB0661
The diurnal hawkmoth Macroglossum stellatarum is
known to feed from a variety of flower species of almost all
colours, forms and sizes. A newly eclosed imago, however,
has to find its first flower by means of an innate flower
template. This study investigates which visual flower
features are represented in this template and their relative
importance. Newly eclosed imagines were tested for their
innate preferences, using artificial flowers made out of
coloured paper or projected onto a screen through
interference filters. The moths were found to have a strong
preference for 440 nm and a weaker preference for 540 nm.
The attractiveness of a colour increases with light intensity.
The background colour, as well as the spectral composition
of the ambient illumination, influences the choice
behaviour. Blue paper disks against a yellowish
background are chosen much more often than the same
disks against a bluish background. Similarly, under
ultraviolet-rich illumination, the preference for 540 nm is
much more pronounced than under yellowish illumination.
Disks of approximately 32 mm in diameter are preferred to
smaller and larger ones, and a sectored pattern is more
attractive than a ring pattern. Pattern preferences are less
pronounced with coloured than with black-and-white
patterns. Tests using combinations of two parameters
reveal that size is more important than colour and that
colour is more important than pattern.
Key words: Macroglossum stellatarum, hawkmoth, Sphingidae,
Lepidoptera, spontaneous choices, innate behaviour, colour vision,
pattern vision.
Summary
Introduction
INNATE PREFERENCES FOR FLOWER FEATURES IN THE HAWKMOTH
MACROGLOSSUM STELLATARUM
ALMUT KELBER*
Lehrstuhl für Biokybernetik, Auf der Morgenstelle 28, D-72076 Tübingen, Germany
Accepted 29 November 1996
*Present address: Centre for Visual Sciences, Research School of Biological Sciences, Australian National University, PO Box 475,
Canberra, ACT 2601, Australia (e-mail: Almut.Kelber@rsbs-central.anu.edu.au).