FISHERY BULLETIN: VOL. 79, NO 3 



radials. In the single osteological preparation of 

 Chaunax examined (Figure 40D), there are three 

 separate, relatively long radials, but the ventral- 

 most element appears to be the result of fusion of 

 three, perhaps indicating the presence of a total of 

 five radials. In Dibranchus (Figure 40E) there are 

 three, relatively short radials, the dorsalmost two 

 lying side-by -side and fused to one another at their 

 proximal and distal ends. 



Skin spines. — Numerous, close-set dermal spines 

 cover the head and body of Antennarius, Lophich- 

 thys, Brachionichthys , and Chaunax; the spines 

 are bifurcated in Antennarius, but simple in 

 Lophichthys, Brachionichthys , and Chaunax. 

 Dermal spines are absent in Tetrabrachium , 

 except for the occasional presence of a spinule 

 associated with an individual pore of the acous- 

 tico-lateralis system. The head and body of Di- 

 branchus are nearly totally enclosed in a cover- 

 ing of thick, nonoverlapping tubercles (Bradbury 

 1967:404). 



PHYLOGENETIC RELATIONSHIPS 



The order Lophiiformes is an assemblage of 18 

 families, 59 genera, and approximately 255 living 

 species of marine teleosts, the monophyletic origin 

 of which seems certain based on the following list 

 of synapomorphic features: 



1) Spinous dorsal fin primitively of six spines, 

 the anteriormost three of which are cephalic 

 in position and modified to serve as a luring 

 apparatus (involving numerous associated 

 specializations, e.g., a medial depression of 

 the anterior portion of the cranium, loss of the 

 nasal bones [nasal of Rosen and Patterson 

 1969 = lateral ethmoid] and supraoccipital 

 lateral-line commissure, and modifications of 

 associated musculature and innervation); 



2) Epiotics separated from parietals and meet- 

 ing on the midline posterior to the supra- 

 occipital; 



3) Gill opening restricted to a small, elongate 

 tubelike opening situated immediately dorsal 

 to, posterior to, or ventral to (rarely partly 

 anterior to) pectoral fin base; 



4) Second ural centrum fused with the first ural 

 and first preural centra to form a single 

 hypural plate (sometimes deeply notched 

 posteriorly) that emanates from a single, 



complex half-centrum (Rosen and Patterson 

 1969:441, text fig. 4E, 60); 



5) Pectoral radials narrow and elongate, the 

 ventralmost radial considerably expanded 

 distally; 



6t Eggs spawned in a double, scroll-shaped mu- 

 cous sheath (Rasquin 1958). 



Since Regan ( 1912), three major lophiiform taxa 

 of equal rank have been recognized by nearly all 

 authors. These taxa, together with their currently 

 recognized families (the 11 families of the bathy- 

 pelagic Ceratioidei excluded), are: 



Suborder Lophioidei 



Family Lophiidae 

 Suborder Antennarioidei 



Family Antennariidae 



Family Brachionichthyidae 



Family Chaunacidae 



Family Ogcocephalidae 

 Suborder Ceratioidei 



In attempting to place Tetrabrachium within 

 the framework of this classification it became 

 apparent that not all of the relationships ex- 

 pressed can be supported by an adherence to 

 cladistic methodology. Although never questioned 

 by any subsequent author, the monophyly of each 

 of Regan's ( 1912) three major lophiiform taxa has 

 not been established. Serious problems lie within 

 the Antennarioidei: a number of synapomorphic 

 features support a sister-group relationship be- 

 tween the Antennariidae and Brachionichthyidae 

 (see below), and between the Chaunacidae and 

 Ogcocephalidae, but no convincing synapomorphy 

 is known at the present time that will link these 

 two larger subgroups. Thus, the problems of inter- 

 preting the interrelationships of higher taxonomic 

 categories within the Antennarioidei, and the 

 relationships of this suborder to the Lophioidei 

 and Ceratioidei are postponed. The following dis- 

 cussion is limited for the most part to Tetrabrach- 

 ium and its relationship to the Antennariidae, 

 to Lophichthys (here given familial rank as 

 suggested by Boeseman 1964), and to the Brachio- 

 nichthyidae. Synapomorphic features that estab- 

 lish monophyly for a group containing the Chau- 

 nacidae and Ogcocephalidae are also enumerated. 

 The relative primitiveness of the character states 

 utilized below was determined by examining their 

 distribution among all available lophiiform ma- 

 terial (47 of the 59 currently recognized genera; 



412 



