Fruit Ripening Signals and Cues in a Madagascan Dry Forest: Haptic Indicators Reliably Indicate Fruit Ripeness to Dichromatic Lemurs

Fruit ripeness can be indicated through changes in chromaticity, luminance, odor, hardness, and size to attract seed dispersing animals. We quantified these attributes for both ripe and unripe fruits of 31 lemur-dispersed plant species in Ankarafantsika National Park, a tropical dry forest in northwestern Madagascar. We used spectroscopy, gas-chromatography mass-spectrometry, and a modified force gauge to quantify chromaticity, luminance, odor, and hardness. We compared these traits between unripe and ripe fruits of each species to determine which traits reliably indicate fruit ripeness across species. Overall, ripe fruits were significantly heavier and softer than unripe fruits. Ripe fruits were not more chromatically-conspicuous or odiferous relative to unripe fruits, nor were ripe fruits more conspicuous in the luminance channel. Contrary to expectation, our findings indicate that, in this particular system, plant-lemur interactions may be strongly mediated by haptic traits, such as fruit hardness, which are consistent and reliable indicators of fruit ripeness. Despite the potential importance of haptic indicators of fruit ripeness, they are underrepresented in the literature on sensory ecology.