Are Archeobatrachian Anurans Ambidextrous? Assessing Handedness in Bombina orientalis

Lateralized behaviors (i.e., handedness), in the absence of conspicuous morphological asymmetry, have been noted in a variety of nonmammalian vertebrates, including certain fishes, frogs, and birds (reviewed in Bisazza et al., 1998; Vallortigara et al., 1999). Lateralized behaviors, such as turning preference during the initiation of a startle response, may facilitate quick reaction. When these behaviors first evolved, though, is unknown. Yamashita et al. (2000) suggested that lateralized behaviors might be common among anurans. However, their presence in any archeobatrachian anurans has not been assessed. One of the most basal archeobatrachian families is the Bombinatoridae (Cannatella, 1999). Here, we test tadpoles and postmetamorphic juveniles of the bombinatorid taxon Bombina orientalis for turning bias and forelimb use, respectively, to determine whether lateralized behaviors occur in this family. There is a fairly extensive literature on handedness in postmetamorphic anurans, at least for neobatrachian taxa. Dill (1977), for example, examined jumping direction in the Pacific tree frog, Hyla regilla, and found a significant left side preference in certain individuals. Bisazza et al. (1997) and Robins et al. (1998) examined handedness in toads of the genus Bufo using a series of protocols. Their first test required each toad to remove a plastic balloon from its head. In this test, Bufo bufo showed significantly greater use of the right forelimb than the left at the population level. Their second test consisted of placing a thin wet strip of paper on a toad's snout and noting which limb it used to clean the paper off. Again, B. bufo showed significant use of the right forelimb. The third test was an underwater righting test in which each toad was submerged in water in a supine position and allowed to right itself. In this test, Bufo marinus exhibited a right forelimb preference, causing it to roll to the left side. In addition, Robins et al. (1998) found lateralized hind limb use by B. marinus, Bufo viridis, and B. bufo when allowed to right themselves on a dry surface. Specifically B. marinus and B. bufo showed a significant preferential use of the right hind limb, and B. viridis showed significant use of the left hind limb. More recently, behavioral laterality was reported in the preferred turning direction of anuran larvae (Wassersug et al., 1999; Yamashita et al., 2000). When tadpoles are startled, or when they descend after surfacing to take a gulp of air, they can turn preferentially

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