Analysis of bumblebee visitation sequences within single bouts: implication of the overstrike effect on short-term memory

Abstract

Pollinators whose foraging habitats consist of several plant types (species or morph) may continue to choose the plant type last visited because information about the type of plant last visited dominates over all other memory contents, in particular of short-term memory. In this study, I extracted this overstrike effect on the plant choices of pollinators by analyzing patterns of visitation sequences within a single round-trip between the hive and foraging patch (bout). First, I simulated the visitation sequences within single bouts with a model to show how factors, including the bees’ plant-type preferences, the arrangement of plants and the effect of overstrike on short-term memory, affect visitation sequences. Here, bees are assumed to forage in a patch consisting of two plant types (H and L). The model predicts that only the effect of overstrike on short-term memory causes assorted visitation sequences according to plant type (within-bout flower constancy). That is, if the overstrike-effect on short-term memory is the primary determinant of plant choice, then bees will fly to a type-L plant after visiting a type-L plant even if they predominantly visit type-H plants and vice versa. Next, I investigated individual bumblebees’ visitation sequences at a patch of artificial inflorescences with a set-up similar to that assumed in the model. Two types of inflorescences were arranged on a Cartesian grid. Assorted visitation sequences according to inflorescence type were observed, depending on the distances among inflorescences. This result supports the hypotheses that bees fly to the same plant type as that last visited because short-term memory is displaced (overstruck) with information about the most recently visited plant type.

Appendix

Possible range of C _p

If N _H −N _L >1 or N _L −N _H > 1, constant flights occur at least once. More generally, the number of constant flights (N _HH +N _LL ) is equal to or larger than |N _H −N _L |−1. Also, N _HH +N _LL is equal to or smaller than the total flight (N _XX’ ). Thus,

$$N_{{X{X}'}} \geqslant N_{{HH}} + N_{{LL}} \geqslant {\left| {N_{H} - N_{L} } \right|} - 1 $$

(A1)

Because N _XX’ =N _X −1, eqn. A1 can be transformed as

$$1 \geqslant \frac{{N_{{HH}} + N_{{LL}} }}{{N_{{X{X}'}} }} \geqslant \frac{{{\left| {N_{H} - N_{L} } \right|} - 1}}{{N_{X} - 1}} $$

(A2)

If N _X is large,

$$ \frac{{{\left| {N_{H} - N_{L} } \right|} - 1}}{{N_{X} - 1}} \cong \frac{{{\left| {N_{H} - N_{L} } \right|}}}{{N_{X} }} = {\left| {F_{H} - F_{L} } \right|} $$

(A3)

Accordingly, if N _X is large, the possible range of C _p (see eqn. 3 in the text) becomes

$$1 \geqslant C_{P} \geqslant {\left| {F_{H} - F_{L} } \right|} $$

(A4)