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Mark Goldy-Brown 11/5/2013 ENT 497A

Lizards, the little known pollinators

Lizards can be pollinators?
When one thinks of pollination, generally an image of a honeybee pops into a persons minds. Perhaps beyond the bee, people will think of butterflies and maybe a few people will manage to remember that hummingbirds and even bats can be pollinators. Insects, birds and mammals have traditionally been thought of as the only pollinators. However, in the last 30 years or so, pollination scientists have realized that another group of animals should be added to this list: lizards. Now why have scientists only recently found out about lizard pollination, you may ask? Well, it is a fairly uncommon phenomenon, occurring mostly on islands and only among herbivorous lizards, which make up only about 1% of all lizard species (Cooper and Vitt, 2002). In fact, lizard pollination seems to be an almost exclusively island phenomenon, largely due to the large environmental differences between island and mainland environments. One group surmises that lizard pollination occurs on islands because lizard densities tend to be higher and lizard predators tend to be less abundant than on the mainland (Olesen and Valido, 2003). This combination of factors lead to diet expansion, meaning in order to survive some lizard species expanded their niche by starting to consume nectar, fruit and even pollen. Over time, what once was just a food source became a plant-pollinator relationship.

The Early Days of Lizard Pollination Research

Pollination in lizards was first studied in the Balearic Islands and in New Zealand, with the earliest studies in New Zealand showing that pollen grains from flowers visited by a native lizard species could stick to the mouth, throat and even belly scales of a lizard (Whitaker, 1987). In fact, they found that in the lizard genus Hoplodactylus, the throat scales were structurally different from those of the rest of the body, which the author proposed could be an adaptation to holding pollen (Whitaker, 1987). The recent focus of lizard pollination however has seemed to shift to the day geckos of the genus Phelsuma, which are found all over the islands of the Indian Ocean. Day geckos tend to be very brightly colored and consume both insects and floral resources such as nectar. One specific species of day gecko which has received particular attention is Phelsuma ornata, or the Mauritius ornate day gecko (Figure 1). Most research on this lizard has occurred in Mauritius, and its surrounding

islands, which are located off the coast of Madagascar in the Indian Ocean. Early research with P. ornata on the Mauritian island of Ile Aux Aigrettes found that these lizards preferred flowering inflorescences rather than fruiting ones and pollen collection tests showed that 86% of the lizards they captured had pollen grains on their throat, lips or belly scales. Due to the documented floral visitations, the distances individual lizards can travel with pollen still stuck to them and the decline of native bird pollinators, they concluded that P. ornata and other Phelsuma spp. were important to promoting outcrossing of many of the native plants on the island (Nyhagen et al., 2001)

Recent Research in Lizard Pollination

More recently however, scientists have begun to further qualify the exact relationship between Phelsuma lizards and native plants. One of the long held mysteries of pollination was the existence of floral nectar color. In general, nectar is clear, however some flowers produce nectar of different colors and this has baffled many scientists. One group of scientists claims that, at least in the Mauritius island area, the nectar coloration serves as a visual cue to pollinators, specifically the Phelsuma spp (Hansen, Beer, Mller, 2006). The Mauritius islands have three flower species that contain colored nectar; Nesocodon mauritianus, which has blood red nectar, Trachetia boutoniana which has red nectar and Trochetia blackburniana, which has yellow nectar (Figure 2).

To test this theory, the team of scientists created fake flowers out of plastic tubes, tape and cardboard. The cardboard was painted various colors and cut out to make the petals, and the tubes were filled with sugar water, to represent nectar. To test if colored nectar made flowers more desirable, a preference choice was set up. Two fake flowers were placed adjacent to each other on a tree, with one flower containing clear nectar while the other had colored nectar, either red or yellow. Interestingly, the population of geckos used in this study was nave to nectar color, meaning that they had never seen this trait before. However, populations of P. ornata on other surrounding islands cooccur with and are known to pollinate the three flowers discussed previously. A nave population was

used because it was easier to work with and also could shed light as to whether any trends discovered would be innate rather than learned. What they found was quite fascinating. The P. ornata had a clear preference for the red and yellow sugar solutions over the clear sugar solutions (Hansen, Beer, Mller, 2006; Figure 3). Surprisingly, the geckos did not prefer the combination of red flower and red sugar water anymore than they did red flower and clear sugar water. This suggested that perhaps the contrast between the nectar color and the floral color is most important to pollinators, rather than just the color of the nectar itself. Given the extreme coloration of many Phelsuma spp. and the apparent innate preferences to colored nectar, it seems reasonable that these lizards would respond well to visual cues, especially relating to color. While an interesting result, it leaves much to be desired. This study was only one small step into the realm of lizard pollination and provides only evidence that there is more to explore in this system. For example, they could not determine whether the colored nectar evolved due to the strong relationship between these flowers and lizard pollinators or whether colored nectar arose due to another pollinator interaction, such as with birds, and the lizards are just taking advantage of it. Despite the limitations of this study, it clearly has opened many new avenues of exploration in the lizard-flower pollination system.

References: Cooper WE and Vitt LJ (2002) Distribution, extent and evolution of plant consumption by lizards. J. Zool. 257: 487-517. Hansen DM, Beer K, and Mller CB (2006) Mauritian coloured nectar no longer a mystery: a visual signal for lizard pollinators. Biol. Lett. 2: 165-168. Nyhagen DF, Kragelund C, Olesen JM, and Jones CG (2001) Insular interactions between lizards and flowers: flower visitation by an endemic Mauritian gecko. J. Trop. Ecol. 17: 755-761. Olesen JM, Valido A (2003) Lizards as pollinators and seed dispersers: an island phenomenon. TRENDS in Ecology and Evolution 18: 177-181. Whitaker AH (1987) The roles of lizards in New Zealand plant reproductive strategies. New Zealand J. Bot. 25: 315-328.