five times, except during the training session for the second part of experiment 2, in which each butterfly was allowed to visit only once see below), we temporarily removed that butterfly from the experimental location until the end of the experiment, the part or the session. When a butterfly visited any flower for a set number of times (i.e. In all behavioural experiments, 10-40 butterflies of each species or each group within a species (see below) were released at the same time and allowed to visit the artificial flowers. We defined a ‘visit’ as a positive response when a butterfly landed and extended its proboscis toward the coloured paper. Together, these studies indicate that flower-visiting insects are generally good learners. Moreover, honeybees and hawkmoths are able to remember unrewarding patterns or colours and avoid them ( Kelber, 1996 Srinivasan et al., 1994). Kandori and Ohsaki, 1996 Laverty, 1980 Lewis, 1986 Raine and Chittka, 2007). With increased experience, bees and butterflies can improve their flower handling skills and/or shorten the time required to find a nectar or pollen source in flowers (e.g. Fan and Hansson, 2001 Goulson and Cory, 1993 Kelber, 1996 Weiss, 1997). For example, butterflies and moths can rapidly undergo reverse conditioning when rewarding and unrewarding flower colours or odours are exchanged (e.g. Kandori and Ohsaki, 1996 Kandori and Ohsaki, 1998 Kinoshita et al., 1999 Lewis, 1986 Lewis, 1989 Swihart and Swihart, 1970) and moths (e.g. Sato and Takasu, 2000 Shafir, 1996 Takasu et al., 2007), hoverflies ( Goulson and Wright, 1998), butterflies (e.g. Dukas and Real, 1991 Heinrich, 1976 Heinrich, 1979 Laverty and Plowright, 1988 Menzel, 1985 Menzel, 1993), wasps (e.g. The ability to learn while foraging for flowers has been demonstrated in many insect taxa, such as bees (e.g. A variety of insects extensively rely on learning for all major life activities, including feeding, predator avoidance, aggregation, social interaction and sexual behaviour ( Dukas, 2008). Dukas and Duan, 2000), and a high learning ability increases fitness ( Raine and Chittka, 2008). The learning abilities of animals are adaptive (e.g. The adaptive significance of superior learning abilities in the larger and longer-lived butterfly species and in females is discussed. This study provides the first evidence that learning abilities related to floral cues differ among butterfly species. Furthermore, female butterflies showed a significantly higher learning rate than males. Among the four species examined, the larger and longer-lived species exhibited higher learning rates. The flower colour learning rates were significantly higher in I. All butterflies that were trained learned the flower colours associated with food. We examined the associative learning of flower colour with nectar in four butterfly species: Idea leuconoe, Argyreus hyperbius, Pieris rapae and Lycaena phlaeas. However, little attention has been focused on differences in floral cue learning abilities among species and sexes. Learning plays an important role in food acquisition for a wide range of insects and has been demonstrated to be essential during flower foraging in taxa such as bees, parasitoid wasps, butterflies and moths.
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