Family Dactylopteridae Cephalacanthidae 303flying gurnards Marine tropical Indo Pacific and Atlantic

Large, blunt, bony head (with spines and keels); body covered with scutelike scales; tremendously enlarged and colorful pectoral fins with inner rays free, total of 28-37 rays; two free spines (the first may be on the nape) before the two dorsal fins; pelvic fins thoracic, each with one spine and four soft rays; no lateral line; 22 vertebrae. Maximum length about 50 cm.

These benthic fishes, which superficially resemble triglids, produce sounds by stridulation by using the hyomandibular bone and "walk" on the sea floor by alternately moving the pelvic fins. The common name arose in the belief that because of their large pectoral fin they could fly or at least glide for short distances. However, they seldom, if ever, leave the substrate and there is no evidence that they ever leave the water and glide.

Two genera, Dactyloptena (Indian and western and central Pacific) and Dactylopterus (Atlantic), with about seven species (e.g., W. N. Eschmeyer in Smith and Heemstra, 1986; Eschmeyer, 1997).

Suborder Scorpaenoidei. Contains the world's most venomous fishes. Usually brightly colored.

In splitting the present family Scorpaenidae, Ishida (1994) recognized the following families in the suborder Scorpaenoidei (also listed in Imamura and Shinohara, 1996): Sebastidae, Setarchidae, Neosebastidae (the latter two being sister taxa), Scorpaenidae, Apistidae, Tetrarogidae (the latter two being sister taxa), Synanceiidae, Congiopodidae (the latter two being sister taxa), Gnathanacanthidae, Aploactinidae, and Pataecidae (the latter two being sister taxa), with the Caracanthidae, of unknown relationships, not being included. Imamura (2004), in his new classification of his superfamily Scorpaenoidea, included the following 20 families (and suggested more might be recognized): Sebastidae (paraphyletic), Sebastolobidae (includes only Sebastolobus), Scorpaenidae (included the Pteroini, Setarchidae, and Trachyscorpia but did not recognize subfamilies or tribes), Apistidae, Tetrarogidae, Synanceiidae (with four subfamilies), Aploactinidae (with three subfamilies), Congiopodidae, Gnathanacanthidae, Pataecidae, Caracanthidae, Eschmeyeridae, Neosebastidae, Plectrogenidae, Parabembridae, Bembridae, Triglidae, Peristediidae, Hoplichthyidae, and Platycephalidae. It should be noted that this includes families placed in the suborder Platycephaloidei. He compared his cladistic results with those of others (e.g., Mandrytsa, 2001), but did not have all families given above included in his study and therefore could provide no independent confirmation of the conclusions of others (this is one reason why I do not split the present family Scorpaenidae).

While I fully acknowledge that the present arrangement, somewhat changed from Nelson (1994), is not satisfactory, I prefer not to recognize numerous additional families (by, for example, raising existing subfamilies to family status), where workers disagree on relationships, until other broad-based studies are done using all genera. The studies noted, in making important contributions, show just how poorly we understand relationships. Users intent on recognizing additional families (as, for example, in splitting the Scorpaenidae)

are free to choose between alternative proposals; I prefer to await more research. While systematic researchers should present the implications of their phylogenetic conclusions to classification, my general advice is that for general use, such as for field guides, when conflicting information exists, it is better to not follow some system that may prove unstable and soon change. On the other hand, all workers in comparative biology will want to be aware of various phylogenetic hypotheses and consult the original research.

Six families with about 82 genera and 473 species.

Family SCORPAENIDAE (304)—scorpionfishes (rockfishes). Marine (rarely in freshwater); all tropical and temperate seas.

Body compressed; head usually with ridges and spines, one or two opercular spines (usually two divergent) and three to five preopercular spines (usually five); suborbital stay usually securely fastened to preopercle (no attachment in some); scales, when present, usually ctenoid; dorsal fin usually single (often with a notch), usually with 11-17 spines and 8-17 soft rays; anal fin with 1-3 spines (usually three) and 3-9 soft rays (usually five); pelvic fin with one spine and 2-5 soft rays (usually five); pectoral fin well developed (11-25 rays), rarely with one to three free lower rays; gill membranes free from isthmus; swim bladder absent in some (e.g., Plectrogenium and Sebastolobus); vertebrae 24-31. Venom gland in dorsal, anal, and pelvic spines. Most have internal fertilization, and some give birth to live young (e.g., Sebastes). Some lay eggs on a gelatinous balloon, and Scorpaena guttata is reported to have an egg balloon that may be as much as 20 cm in diameter. Many species are commercially important.

At least 56 genera with about 418 species (e.g., Poss and Eschmeyer, 2003). Most species are in the Indian and Pacific oceans. For comments on the classification see above under suborder Scorpaenoidei. The sequencing of the following subfamilies, recognized as families from some workers as noted above, is based, in part, on conclusions in some of the works listed above. Extensive work over many decades has been done on this family by W. N. Eschmeyer, with much work also by S. G. Poss and more recently by H. Motomura.

Subfamily Sebastinae. Imamura (2004) recognized Trachyscorpia with the Scorpaenini (as here used) and placed Adelosebastes in his paraphyletic Sebastidae (as he states, it would be cladistically reasonable to have each genus treated as a separate family, something he and certainly not I were prepared to do), leaving only Sebastolobus in his Sebastolobidae.

Seven genera and about 133 species. Extensive information is given on the members of this subfamily, known as rockfishes, in the northeast Pacific by Love et al. (2002).

tribe sebastinae. Four genera, Helicolenus, Hozukius, Sebastes, and Sebastiscus, with about 128 species. Helicolenus and Sebastes occur in all oceans, whereas Sebastiscus and Hozukius occur only in the western Pacific. Kai et al. (2003)

suggested that Helicolenus, Hozukius are more closely related to Sebastes than to Sebastiscus. The live-bearing genus Sebastes is the largest in the family with about 110 species (almost all of them occurring in the North Pacific).

tribe sebastolobinae. Three genera, Adelosebastes, Sebastolobus (with 15-17 dorsal spines, highest for the family), and Trachyscorpia, with five species.

Subfamily Setarchinae. Three genera, Ectreposebastes, Lioscorpius, and Setarches, with five species. Imamura (2004) recognized this subfamily with the Scorpaenini (as here used).

Subfamily Neosebastinae. Two genera, Maxillicosta (5) and Neosebastes (12), with 17 species (Motomura, 2004a). As noted by Motomura (2004a), Ishida (1994) inferred that his families Neosebastidae and Setarchidae had a sister relationship and were secondarily divided from other scorpaenids. Imamura (1996), how-ever, suggested that Setarches was more closely related to Pontinus and Scorpaena (tribe Scorpaenini herein), Neosebastes being sister to a clade comprising a species of Tetraroginae, two genera of Synanceiinae, an Aploactinidae, and an Apistinae. Smith and Wheeler (2004), in a molecular analysis, showed that Maxillicosta and Congiopodus had a sister relationship. As Motomura (2004a) concluded, the systematic position and relationships of the family Neosebastidae (as he recognized it), still lacks an established basis. Imamura (2004) regarded this subfamily (at the family level) as sister to the Platycephaloidei (as given here) and further studies await with interest.

Subfamily Scorpaeninae. At least 20 genera and about 185 species.

tribe scorpaenini

At least 15 genera (e.g., Idiastion, Iracundus, Neomerinthe, Parascorpaena, Phenacoscorpius, Pontinus, Pteroidichthys, Pteropelor, Rhinopias, Scorpaena, Scorpaenodes, Scorpaenopsis, Sebastapistes, and Taenianotus) with about 165 species (e.g., Randall and Eschmeyer, 2002; Randall and Greenfield, 2004).

tribe pteroini. Five genera, Brachypterois, Dendrochirus, Ebosia, Parapterois, and Pterois (highly venomous lionfishes and turkeyfishes), with about 20 species.

Subfamily Apistinae. Three monotypic genera, Apistops, Apistus, and Cheroscorpaena.. Members of this taxon have one or three free lower pectoral rays and a bilobed swim bladder.

Subfamily Tetraroginae (sailback scorpionfishes or wasp fishes)

At least 11 genera (e.g., Ablabys, Amblyapistus, Centropogon,, Cottapistus, Neocentropogon, Notesthes, Ocosia, Paracentropogon,, Tetraroge, and Vespicula) with about 38 species. Tetrarogines are extremely venomous. They show some resemblance to the Aploactinidae. Notesthes robusta of coastal eastern Australia is primarily a freshwater fish.

Subfamily Synanceinae. Body scaleless (except for buried scales along the lateral line and other parts of the body), usually covered with skin glands; head large; swim bladder usually absent; venom glands present near base of hypodermiclike dorsal fin spines. The neurotoxin of these fishes is the most deadly of the fish venoms and can be fatal to humans. The fish is particularly dangerous because it usually rests in a half-buried position, looking much like a rock.

About nine genera and about 35 species.

tribe minoini. Lowermost ray of pectoral fin separated from the other 11 rays, fitted at its tip with a peculiar "cap"; body smooth; dorsal fin with 8-12 spines and 10-14 soft rays (4 spines and 18 soft rays in one species); anal fin with two spines and 7-11 soft rays; pelvic fin with one spine and five soft rays; soft fin rays unbranched; swim bladder present or absent; vertebrae 24-27. Maximum length usually 15 cm. Members of this group occur on mud and sand bottoms from about 10-420 m in the western Pacific and Indian oceans. They are thought to use the free pectoral ray for "walking" on the bottom.

The Fishes Indian Pacific

One genus, Minous, with 11 species.

tribe choridactylini. Two (Inimicus) or three (Choridactylus) lowermost pectoral rays separated from rest; body often with warts or lumps (caused by buried scales); dorsal fin with 12-18 spines and 5-10 soft rays; anal fin with two spines and 8-13 soft rays; pelvic fin with one spine and five soft rays; most soft fin rays branched; vertebrae 26-30. Members of this group occur on sand and silty bottoms from near shore to about 90 m in the western Pacific and Indian oceans. Imamura (2004) placed the two genera in separate subfamilies.

Two genera, Inimicus with 10 species and Choridactylus with four species (e.g., Poss and Mee, 1995).

tribe synanceini (stonefishes) . No free pectoral rays; skin glands present (appearing as "warts" in most species) and usually scattered over the body; dorsal fin with 11-17 spines and 4-14 soft rays; anal fin with 2-4 spines and 4-14 soft rays; pelvic fin with one spine and 3-5 soft rays; pectoral fin rays 11-19; vertebrae 23-30.

Two species, Erosa erosa (Japan to Australia) and Dampierosa daruma (northwestern Australia), have a terminal mouth that is slightly oblique and lateral eyes that are directed outward. The remaining species have a vertical or superior mouth and dorsal eyes that are directed outward and upward or only upward. Some species are known from rivers.

Dampierosa Daruma

Six genera, the monotypic Erosa, Dampierosa, Pseudosynanceia, Leptosynanceia, and Trachicephalus and Synanceia (with five species), with a total of 10 species.

Subfamily Plectrogeninae. One genus, Plectrogenium, with two species. There is good evidence showing a relationship with the Platycephaloidei (Imamura, 1996, 2004).

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