Scales absent; body eel-like; mouth small, lower jaw protruding in front of upper; dorsal and anal fins confluent with caudal fin, dorsal fin with 37-50 rays, caudal fin with 4-6 rays, and anal fin with 34-43 rays; dorsal fin origin posterior, well behind pectoral fin; two nostrils on each side of head; no sensory pores on head (neuromasts free); pelvic fins absent; pectoral fin small, with 6-8 rays; ovaries bilobed; vertebrae 54-73. Maximum length about 6 cm.
Nelson (1994) noted reasons for placing and for not placing this family near the zoarcids or, as here, the aphyonids. Nielsen et al. (1999) did not regard it as an ophidiiform. Clearly, it is a family without a home. For purposes of this classification, I leave it where placed in Nelson (1994). Just as then, more research is desirable to determine if parabrotulids are aphyonid derivatives, related to the zoarcids, or related to some other taxa.
Two genera, Parabrotula and Leucobrotula, with three species (Miya and Nielsen, 1991).
Pediculati. Patterson and Rosen (1989) considered the batrachoidiforms and lophiiforms to be sister groups and applied the term Pediculati to include both orders. There is doubt, based on molecular biology, that this group is monophyletic (see above under Paracanthopterygii).
Order BATRACHOIDIFORMES (Haplodoci) (46)—toadfishes. Body usually scaleless (small cycloid scales in some); head large with eyes more dorsal than lateral; mouth large and bordered by premaxilla and maxilla; pore (foramen) in axil of pectoral fin in some; pelvic fins jugular (in front of pectorals), with one spine and two or three soft rays; three pairs of gills; gill membrane broadlyjoined to isthmus; branchiostegal rays six; four or five pectoral radials; swim bladder present; upper hypurals with peculiar intervertebral-like basal articulation with rest of caudal skeleton; no ribs, epiotics, or intercalars; no pyloric caeca.
Some members can produce audible sounds with the swim bladder and can live out of water for several hours. Most are drab colored.
Family BATRACHOIDIDAE (226)—toadfishes. Marine (primarily coastal benthic; rarely entering brackish water, a few species confined to freshwater); Atlantic, Indian, and Pacific.
Three subfamilies with 22 genera and 78 species.
Subfamily Batrachoidinae. Off coasts of the Americas, Africa, Europe, southern Asia, and Australia.
Three solid dorsal spines and solid opercular spine, no venom glands; sub-opercular spines present; body with or without scales (cycloid); no photophores; axillary gland at pectoral base present or absent; canine teeth absent; usually one or three lateral lines.
Toadfishes generally occur on sand and mud bottoms, although species of Sanopus occur in coral reefs.
Eighteen genera, Allenbatrachus, Amphichthys, Austrobatrachus, Barchatus, Batrachoides,, Batrachomoeus, Batrichthys,, Bifax, Chatrabus, Halobatrachus, Halophryne, Opsanus, Perulibatrachus (a replacement name for the fossil Parabatrachus), Potamobatrachus, Riekertia,, Sanopus, Tharbacus, and Triathalassothia, with about 52 species (e.g., Collette, 1995a, 2001, 2003a; Greenfield, 1996, 1998; Greenfield et al., 1994; Greenfield and Smith, 2004).
Subfamily Porichthyinae. Eastern Pacific and western Atlantic.
Two solid dorsal spines and solid opercular spine, no venom glands; no sub-opercular spines; body scaleless; photophores present or absent; axillary gland absent; canine teeth present; several lateral lines. Two genera with 15 species.
Aphos. Lacks photophores; one southeastern Pacific (Peru and Chile) species (Walker and Rosenblatt, 1988).
Porichthys (midshipmen). Numerous photophores (this is one of the few shallow-water fishes that possess photophores); four lateral lines. Fourteen species, eight along the eastern Pacific (British Columbia to northern Peru and Galapagos Islands) and six along the western Atlantic (Virginia to Argentina, but generally absent from the West Indies) (Walker and Rosenblatt, 1988). Maximum length 43 cm, in P. myriaster.
Subfamily Thalassophryninae. Eastern Pacific and western Atlantic.
Two hollow dorsal spines and hollow opercular spine (serving as a venom-injecting apparatus capable of producing extremely painful wounds, connecting with venom glands; no subopercular spines; body scaleless; no photophores; no canine teeth; lateral line single or absent; pectoral fin rays 13-18.
Two genera with 11 species (e.g., Collette, 1973).
Daector. Second dorsal fin rays 22-33; anal fin rays 21-30; distinct glands with pores between bases of upper 3-7 pectoral fin rays on inner surface of fin (similar to Opsanus); vertebrae 31-40. Four tropical eastern Pacific marine species and D. quadrizonatus from freshwater, Columbia (Atrato basin, Atlantic drainage).
Thalassophryne. Second dorsal fin rays 17-22; anal fin rays 16-20; indistinct glandular tissue lacking pores scattered along upper rays of pectoral fin;
vertebrae 26-30. Five western Atlantic marine species (Panama and South America) and T. amazonica, known only from the Amazon River.
Order LOPHIIFORMES (47)—anglerfishes. First ray of spinous dorsal, if present, on head and transformed into illicium (line) and esca (bait), a device for attracting prey to mouth; pelvic fins, if present, in front of pectorals, with one spine and four (rarely) or five soft rays; gill opening small, tubelike, at or behind (rarely partly in front of) pectoral fin base; five or six branchiostegal rays; no ribs; pectoral radials 2-5, narrow and elongate; first vertebra fused to skull; swim bladder, when present, physoclistous.
The following classification is based on the phylogenetic conclusions of Pietsch and Grobecker (1987). However, in contrast to the three suborders I present, these two workers recognized five suborders comprising the following postulated monophyletic groups.
1. Ogcocephaloidei and Ceratioidei—united by, e.g., second dorsal spine reduced to a small remnant (except well developed in the Diceratiidae), third dorsal spine and pterygiophore absent, and posttemporal fused to cranium.
2. Chaunacoidei—the sister group of the above by sharing second dorsal spine (elongate in the Chaunacidae) embedded beneath the skin of the head, and gill filaments of the first gill arch absent. The three suborders in 1 and 2 are included here in the suborder Ogcocephaloidei.
3. Antennarioidei—the sister group of the above by sharing, e.g., eggs and larvae much smaller than those of lophiids; dorsal fin spines reduced to three or less.
4. The Lophioidei—considered to be the primitive sister group of the above.
Eighteen families with about 66 genera and 313 species. All are marine. Most species occur in deep water.
Suborder Lophioidei. Pelvic fins present; spinous dorsal fin behind head with one to three spines; fourth pharyngobranchial present and toothed; pseudo-branch large; body scaleless; frontals united.
Family LOPHIIDAE (227)—goosefishes. Marine; Arctic, Atlantic (including the Mediterranean Sea), Indian, and Pacific.
Huge, wide, flattened head (head rounded in Sladenia); teeth well developed; fringe of small flaps extending around lower jaw and along sides of head onto body; pectoral fin rays 13-28; second dorsal fin with 8-12 soft rays; anal fin with 6-10 rays; vertebrae 18 or 19 and, in Lophius, 26-31.
The mobile fishing apparatus has a flap of flesh at its tip that acts like a lure, attracting prey within reach of its large mouth. Size up to 1.2 m.
Four genera, Lophiodes (13), Lophiomus (1), Lophius (8), and Sladenia (3), with 25 species (Caruso, 1985). Fossils of Eocene age (e.g., Eosladenia) are known (Bannikov, 2004a).
Suborder Antennarioidei. Spinous dorsal fin consisting of three separate cephalic spines (the first is the modified illicium; the second dorsal spine may be short, but it is never embedded beneath the skin); pterygiophores of illicium and third spine of dorsal fin with highly compressed dorsal expansions.
In contrast to those taxa placed in the superfamily Ceratioidea, members of the two families of this suborder and of the Chaunacidae and Ogcocephalidae share the following features: pelvic fins present; pectorals usually armlike; pseudobranch (small) present; swim bladder present (most Antennariinae only) or absent; body usually covered with small, closely set dermal spines; frontals united posteriorly, but usually separated from each other anteriorly. Most species are benthic; only Histrio is epipelagic, occurring in sargassum.
Four families with 15 genera and 48 species. Unlike in Nelson (1994), Tetrabrachium and Lophichthys are placed in separate families following Pietsch and Grobecker (1987).
Family ANTENNARIIDAE (228)—frogfishes. Marine; all tropical and subtropical seas (absent from the Mediterranean), occasionally temperate (e.g., western Atlantic and southern Australia).
Deep-bodied (globose); nape not conspicuously humped; mouth large; eyes lateral; body covered with loose skin, naked or with denticles; gill opening below or behind base of pectoral fin; pelvic fin of one spine and five soft rays; parietals separated by supraoccipital; pectoral radials 3; swim bladder usually present (absent in Kuiterichthys and Tathicarpus); soft dorsal fin rays 10-16; soft anal fin rays 6-10;
pectoral fin rays 6-14; vertebrae 18-23; palatine teeth present. The fishing pole (illicium) of frogfishes, a modification of the first dorsal spine, is pronounced and highly variable between species. Maximum length 36 cm, some only 3 cm.
Frogfishes are benthic except for the widespread and pelagic Histrio histrio, which uses its prehensile pectoral fin for "clasping" or moving on floating sar-gassum. The Indo-Australian species Antennarius biocellatus is the only frogfish known to occur in brackish and freshwater. Pietsch and Grobecker (1987) give distributional information on the various species. The feeding dynamics of frogfishes are described by Pietsch and Grobecker (1987); they note cases of aggressive mimicry in which the lures of different species resemble a poly-chaete, an amphipod, and a small fish. In addition, there are a few species with parental care in which the eggs may also be acting as a lure.
Twelve genera, Allenichthys, Antennarius, Antennatus, Echinophryne, Histiophryne, Histrio, Kuiterichthys, Lophiocharon, Nudiantennarius, Phyllophryne, Rhycherus, and Tathicarpus, with 42 species (e.g., Pietsch and Grobecker, 1987; Ohnishi et al., 1997; Randall and Holcom, 2001). Over half the species are placed in the genus Antennarius, this genus appears to be the most primitive for the group, but unlike the other genera, its monophyly has not been established.
Family TETRABRACHIIDAE (229)—tetrabrachiid frogfishes. Marine; western and northern coasts of Australia, southern coast of New Guinea, and the south Molucca Islands of Indonesia.
Body elongate and strongly compressed; mouth small; eyes small and dorsal; swim bladder absent; nape humped; soft dorsal fin rays 16 or 17; anal fin rays 11 or 12; pectoral fin rays 9, fin divided into two portions; palatine teeth absent. Maximum length about 7 cm.
One species, Tetrabrachium ocellatum (Pietsch and Grobecker, 1987).
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