COLLETTE ET AL.: BELONIFORMES 



345 



(subgenus Cheilopogon s. str.) are flaplike and fringed (Fig. 1811). 

 Kovalevskaya (1982) considered this to result from the fusion 

 of paired barbels and our examination of Ch. pmnatibarbatus 

 calijorniciis supports this. The barbel is single in Cypselurus 

 (subgenus Cypselurus s. str.) and Exocoetus monocirrhus (Fig. 

 1 8 1 F; Parin, 1961; Kovalevskaya, 1982). Some species of Cyp- 

 selurus (subgenus Poecilocypselurus, Cy. poecilopterus and Cy. 

 starksi) do not develop a barbel, nor do the remaining two 

 species of Exocoetus, E. obtusirostris and E. volilans. 



Melanistic dorsal fin lobe 



Pelagic members of three families (all except Scomberesoci- 

 dae) develop prominent melanistic lobes in the dorsal fin. The 

 lobe is in the posterior part of the dorsal fin in the Belonidae 

 and Hemiramphidae but in the middle of the fin in the Exo- 

 coetidae so presence of the lobe is not necessarily homologous. 



Beionidae.— A blennes and Tylosurus are characterized by hav- 

 ing a prominent enlarged melanistic lobe in the posterior part 

 of the dorsal fin (Fig. 179D, G-J). Other genera of needlefishes 

 (Fig. 1 79) lack any trace of this posterior dorsal lobe. Breder 

 (1932: plates 3-5) illustrated the development of this posterior 

 lobe in T. acus and T. crocodilus and its absence in Strongylura 

 and Platybelone. Parin (1967) left an Australian species difficult 

 to place in either Tylosurus or Strongylura in a monotypic genus 

 described by Whitley, Lhotskia gavialoides. A juvenile with a 

 well-developed posterior dorsal lobe, captured by Collette. con- 

 vinces us that it is a species of Tylosurus (Fig. 1 79H). The lobe 

 is apparently sloughed off in Tylosurus crocodilus (Breder and 

 Rasquin, 1952), resorbedin T. aoM (Breder and Rasquin. 1954), 

 and retained in adult Ablennes. 



Hemiramphidae.— Juveniles of Hemiramphus and Oxypo- 

 rhamphus develop a darkened posterior lobe on the dorsal fin 

 (Fig. 180) similar to that present in two genera of needlefishes, 

 Ablennes and Tylosurus. 



Exocoetidae. — In juveniles of many species oi Cheilopogon. the 

 middle portion of the dorsal fin develops a melanistic lobe (Fig. 

 1 8 1 H). This is reminiscent of the adult stage of Parexocoetus 

 and Eodiator acutus. 



Body bars 



Juveniles of some species in three exocoetoid families (all 

 except Scomberesocidae) have vertical bars on their body. 



Belonidae.— Juveniles of two species of Tylosurus, T. gavi- 

 aloides (Fig. 179H) and T. acus (see Collette and Parin. 1970: 

 fig. 12) and .Ablennes hians have bars. These bars are retamed 

 in adult .-iblennes as is the posterior dorsal fin lobe. 



Hemiramphidae. — The 10 species of the genus Hemiramphus 

 all have a series of broad vertical bars on the body (Fig. 180A- 

 E) at some stage of their development. Body bars are retained 

 for different periods of time during development: all body bars 

 are lost before 105 mm SL in He. lutkei and He. depauperatus 

 (Parin et al., 1980: fig. 32), before 120 mm SL in He. bermu- 

 densis and He. brasiliensis (Collette, 1962: fig. 1), but are re- 

 tained past 175 mm SL in He. balao; one blotch is retained 

 throughout life in He. robustus, and all are retained in He. far. 



Pelvic fin pigment 



All 10 species of Hemiramphus also have pigmented pelvic 

 fins as juveniles (Fig. 183). The patterns of pelvic fin pigmen- 

 tation divide the genus into two species groups, one with pig- 

 mentation concentrated proximally on the fin (balao group. Fig. 

 183, top two rows), the other with pigment absent basally and 

 concentrated distally (J'ar-brasiliensis group. Fig. 183. bottom 

 row). Body bars and pelvic fin pigmentation are absent in Hy- 

 porhamphus. 



Exocoetidae. — In late larval and juvenile stages of many flying- 

 fishes, Exocoetus, Cheilopogon (at least some species in all sub- 

 genera except possibly Paracypselurus, for which we lack data), 

 Cypselurus (subgenus Poecilocypselurus— see Imai, 1959), and 

 Hirundichthys oxycephalus (Imai, 1960) transverse stripes de- 

 velop on the abdomen and sides of the body which disappear 

 (sometimes leaving traces) in adults. The coloration of the larvae 

 and particularly of the juveniles of flyingfishes is diverse, and, 

 as a rule, differs greatly from the coloration of adults. A partic- 

 ularly bright variegated coloration is characteristic of young of 

 neritic species living among algae (Parin, 1961; Kovalevskaya, 

 1982). 



Relationships 



Beloniformes 



The Beloniformes were defined by 7 characters by Rosen and 

 Parenti (1981:16). Meristic characters for the beloniform genera 

 are summarized in Table 92. A cladogram for the families and 

 higher taxa of the Beloniformes is presented as Fig. 184. 



Adrianichthyoidei 



Rosen and Parenti (1981) defined the adrianichthyoids by 5 

 characters. Larval adrianichthyids also differ from exocoetoids 

 in having a shorter preanal distance, 40-50% of standard length. 

 Rosen and Parenti (1981) included the Horaichthyidae and Ory- 

 ziidae in the Adrianichthyidae. By this definition the Adrianich- 

 thyidae includes four genera, .Adrianichthys, Horaichthys, Ory- 

 :ias and Xenopoecihis with a total of 1 1 species (Nelson, 1976). 

 These fishes inhabit fresh and/or brackish waters from India 

 and Japan to the Indo-Australian Archipelago. 



Fig. 179. Halfbeak stages of Belonidae, arranged by relative length of upper Jaw. (A) Belonion apodion Collelle: USNM 199540; Brazil, Borba; 



29.4 mm BL; (B) Belonion dihranchodon Collette; USNM 199463; Venezuela, Rio Atabapo; 38.2 mm BL; (C) Strongylura marina (Walbaum), 

 USNM 189006; Nicaragua; 23.5 mm BL; (D) Ablennes hians (Valenciennes); USNM 188843; Gulf of Honduras; 36.1 mm BL; (E) Platybelone 

 argatus argalus (Le Sueur) USNM 198102; 39°28'N, 69°30'W; 96 mm BL; (F) Strongylura p.v;fa (Girard); SIO H47-158-23A; Calif, La Jolla; 



72.5 mm BL; (G) Tylosurus acus acus (Ucepede); USNM 1 98402; 38°00'N, 65''25'W; 1 30 mm BL; (H) Tylosurus gavialoides (Castelnau); USNM 

 226666; Australia, New South Wales; 72.5 mm BL; (1) T. choram (Forsskal); USNM 147438; Red Sea; 95.0 mm BL; (J) T. c. crocodilus (Peron 

 and Le Sueur); USNM 198407; 37°08'N, 66°14'W; 96.3 mm BL. A-G, 1-J drawn by Mildred H. Carrington; H by Keiko Hiratsuka Moore; A- 

 C from Collette (1966: tig. 1). 



