Annals of the History and Philosophy of Biology

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70 Annals of the History and Philosophy of Biology, Vol. 10 (2005) Ulrich Kutschera (members of the genera Pelecus and Alburnus) occasionally "walk on the surface of the water" in response to attacks from predators. Halfbeaks (Hemirhamphus, Oxyporhamphus) are intermediate forms that display predator-driven jumps out of the water that are reminiscent of the flights of the exocoetids (Fig. 6 A, B). Hence, the evolution of flight in the Beloniformes originated with now extinct species that temporarily "walked out of the water" to escape predators. Lorenz (1963) did not distinguish between exocoetids that have two versus four "wings" (Fig. 8) However, he pointed out that the surface of the water, viewed from below, looks like a mirror: the aquatic prey organism, driven out of the liquid medium, becomes invisible to the predator. Fig. 7: Oscillatory side-to-side movement of the tail of a flyingfish, viewed from above (A). A fourwinged (biplane-type) cypselurid fish, front view in flight (B) (Adapted from Breder 1930). The evolution of gliding in Beloniform fishes: a synthesis Darwin (1872) proposed that the phylogenetic development of novel body plans is driven by the same mechanisms that cause the origin of new varieties and species. This classical concept of "phylogenetic gradualism" (Gould 2002) has developed into a basic tenet of the modern theory of biological evolution: large phenotypic changes (origin of higher taxa) are brought about by successive microevolutionary processes. Although exceptions to this rule exist, there is consensus among the majority of biologists that macroevolution (phylogenetic development above the species level) is the product of numerous microevolutionary steps (Mayr 1963, 2001; Futuyma 1998; Zimmer 1998, Carroll 2000, 2001; Simons 2002, Kutschera and Niklas 2004, 2005). The predator-driven development of gliding in Beloniform fishes discussed here is an example of a macroevolutionary trend. It is obvious that the beating tail of the exocoetids, which propels the fish clear of the water, and the enlarged pectoral fins are organs that have undergone an "intensification and/or change in function" (Mayr 1963): the tail acts as a "motor", the fins are "wings", the flyingfish displays the aerodynamic properties of an aeroplane or a hirundine bird (swallow).
Predator-driven macroevolution in flyingfishes inferred from behavioural studies Fig. 8: Cladogram of halfbeaks (Hemirhamphidae) and flyingfishes (Exocoetidae). According to this scheme, the shortwing flyingfish Oxyporhamphus is a member of the Exocoetidae. Three four-winged (biplane-type) exocoetids are depicted in the lower part of the cladogram (Cypselurus, Prognichtys, Hirundichthys) (Adapted from Dasilao and Sasaki, 1998). A hypothesis for this macroevolutionary trend in the Beloniformes, based on observations of extant "model organisms" that represent various stages in phylogeny, is depicted in Figure 9. This scheme is an expanded and modified version of the "historical reconstruction" presented by Lorenz (1963), with reference to Klausewitz (1960). The data of Dasilao and Sasaki (1998) (Fig. 8) are largely in accordance with the phylogenetic hypothesis discussed here. Annals of the History and Philosophy of Biology, Vol. 10 (2005) 71
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Deutsche Gesellschaft für Geschich
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Preface A new yearbook for history
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