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72 Annals of the History and Philosophy of Biology, Vol. 10 (2005) Ulrich Kutschera Members of the fish family Cyprinidae (minnows or carps) that inhabit the upper region of the waters (Pelecus, Alburnus and others), are able to "walk on the surface" to escape predatory attacks (Fig. 9 A). These "walking fish" may represent the ancestral stage in this evolutionary trend. Halfbeaks, represented by members of the genus Hemirhamphus, are prone to leap out of the water; they usually perform a short "walk on the surface" before they temporarily leave the liquid medium (Fig. 9 B). According to Lorenz (1963), the hemirhamphid Oxyporhamphus represents an intermediate form between a typical halfbeak and a true flyingfish. This "shortwing flyingfish" (Smith and Heemstra 1986) has an elongated lower jaw only as a juvenile (i.e., it recapitulates the halfbeak stage during ontogeny), a deeply forked caudal fin (lower lobe longer than upper), and longer wing-like pectoral fins than other typical halfbeaks (Fig. 9 C). Dasilao et al. (1997) have provided evidence that, based on morphological data, the halfbeak Oxyporhamphus should be considered a basal flyingfish, as suggested by Lorenz (1963). However, molecular data presented by Lovejoy et al. (2004) place Oxyporhamphus within the Hemirhamphus clade. These contradictory results indicate that the "shortwing-halfbeak" Oxyporhamphus is an extant intermediate form between the Hemiramphidae and the Exocoetidae. Breder (1930) was the first to distinguish between two categories of flyingfishes (Exocoetidae), "two-wingers" (Fodiator, Parexocoetus, Exocoetus etc.) in which the enlarged pectoral fins make up most of the lifting surfaces, and "four-wingers" (Cypselurus, Prognichthys, Hirundichthys etc.) in which both pectoral and pelvic fins are hypertrophied (Fig. 8). According to Collette and Parin (1998) two-winged exocoetids may glide for a distance of 25 m, whereas four-winged species may achieve 200 m or more with the extra lift generated by the enlarged pelvic fins. However, both types of exocoetids use their hypertrophied lower portion of the asymmetrical tail fin to provide the impetus for the free flight (Fig. 7 A). A number of studies have shown that "two-wingers" like Parexocoetus (Fig. 9 D), along with Exocoetus and Fodiator, are the least sophisticated gliders. These "primitive" flyingfishes are at the base of the exocoetid tree, as studied by cladistic methods (Fig. 8; Dasilao and Sasaki 1998). It is obvious that the more sophisticated "biplane gliders" (Cypselurus and related taxa) (Fig. 9 E) evolved from more basal "two-wingers"; these "living airplanes" represent the extant peak in Exocoetid evolution. They build up speed by taxiing like aircraft and resemble herring-like swallows. The subtropical flyingfish Hirundichthys depicted in Fig. 8 is a bird-like vertebrate, with black fins that look like the wings of some Aves that glide over large distances (Smith and Heemstra 1986). Lovejoy et al. (2004) reconstructed the phylogeny of 54 species of Beloniform fishes, using fragments of two mitochondrial and two nuclear genes. These molecular data generally confirm the concept depicted here (Fig. 9), with the exception that the intermediate form Oxyporhamphus occurs deeply within the Hemirhamphus clade.
Predator-driven macroevolution in flyingfishes inferred from behavioural studies Fig. 9: Predator-driven evolution of gliding in Beloniform fishes, based on behavioural studies of extant species. Cyprinid (Pelecus) that occasionally leaps at the surface (A), halfbeak (Hemirhamphus) that jumps out of the water (B). The short-wing flyingfish (Oxyporhamphus), depicted as adult and juvenile individuum, represents an intermediate form that recapitulates the halfbeak-stage during ontogenesis (C). Monoplane-type flyingfish (Parexocoetus) that has a single set of long pectoral fins (wings) (D) and biplane-type (Cypselurus) that has under wings (pelvic fins) staggered far back from the main wings (E) (Adapted from Lorenz 1963). In addition, the phylogenetic trees reconstructed on the basis of DNA-sequence data shed light on the ontogenetic recapitulation of the "halfbeak stage" in the "shortwing flyingfish" Oxyporhamphus and related taxa (Lovejoy 2000, Lovejoy et al. 2004; for his- Annals of the History and Philosophy of Biology, Vol. 10 (2005) 73
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Preface A new yearbook for history
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