WASHINGTON ET AL.: SCORPAENIFORMES 



441 



Table 111. Continued. 



• Last ray single (usual condition is a double ray). 

 " I + + V + I or I + I + V + I = 7-8 spines. 

 **" Placement uncertain. 

 ' Data supplied by Poss. 

 - From Poss (1982). 



nanuni is closely related to the bembrids," and that "it is very 

 probable, therefore, that the platycephalids, bembrids and hop- 

 lichthyids arose from an ancestor not very unlike the scorpaenid. 

 Plntrogeniuin naiiuin." Matsubara and Ochiai (1955) present 

 additional characters which support this view. Other characters 

 observed by one of us (WNE) which support this conclusion 

 include, "similar caudal skeletons and scales" [comparison of 

 bembrid Parabembras ciirtis (SU 49456, cleared and stained) 

 and Pleclrogeiiiu?}!]. At present, this available evidence suggests 

 that the scorpaenid subfamilies Sebastolobinae and Plectrogen- 

 inae and the families Bembridae, Platycephalidae and Hoplich- 

 thyidae may form a monophyletic assemblage. 



Another scorpaenid subfamily, the Apistinae, also has ques- 

 tionable relationships within the family. Matsubara ( 1 943) placed 

 Apislus at the base of his Cocolropiis stem which led to a number 

 of specialized scorpaenid groups. However, the Apistinae have 

 a "unique", bilobed gasbladder with an intrinsic muscle, unlike 

 other scorpaenids (Matsubara, 1943; Hallacher, 1974). The trig- 

 lids and peristediids possess a similar gasbladder. Other char- 

 acters which appear to unite the Apistinae, Triglidae, and Per- 

 istediidae include elongate pectoral fin rays, 1 to 3 lower free 

 pectoral fin rays(l in Apistus. 3 in Chcroscorpaena. 3 in triglids, 

 and 2 in peristediids) and shape and expansion of the head bones 

 (especially the infraorbidal bones). These characters suggest that 

 the scaled, less bony-headed Apistinae may be the primitive 

 sister group of the Triglidae and Penstediidae. This would in- 

 volve the independent development of a moveable, preorbital 

 bone with long spine in the Apistinae. If in fact, the Apistinae 

 forms part of a monophyletic assemblage with the triglids and 

 peristediids, a change in classification would be warranted. 



Matsubara (1943) recognized five Japanese genera within his 

 subfamily Congiopinae (sic) which more recently have been 

 placed in the tentative scorpaenid subfamily, Tetraroginae (Poss 

 and Eschmeyer. 1975; see also Smith, 1958b). The presently 

 recognized Congiopodidae is considered to contain 7 to 9 species, 

 all of which are confined to the Southern Hemisphere (More- 

 land, 1960; Hureau, 1971). Moreland (1960:241) slated: "the 

 Congiopodidae show relationship with the Scorpaenidae, par- 

 ticularly with Snyderina and Ocosia from Japan [studied by 

 Matsubara (1943)], and are clearly derived from a scorpaenid 

 stock of perhaps Indo-Pacific origin." 



We tentatively include the Dactylopteridae in our discussion 

 of the Scorpaenoidei, however relationships of these fishes re- 

 main uncertain. They have been variously placed in their own 

 order (Regan, 1913a; Berg, 1940; Greenwood et al., 1966; Lau- 

 der and Liem, 1983) and as a suborder of the Scorpaeniformes 

 (Gill, 1888; Nelson, 1976). Many workers have noted that the 

 dactylopterids differ markedly from scorpaeniforms in a number 

 of osteological characters such as: 1) nasals fused into a median 

 plate; 2) very large extrascapulars; and, 3) mesethmoid and in- 

 tercalar absent. 



Matsubara (1943) suggested that despite these notable differ- 

 ences, the dactylopterids possess the characteristic suborbital 

 arrangement of bones of the generalized scorpaenids and triglids. 

 .•Accordingly, Matsubara (1943) placed them near the triglids 

 and peristediids, evolving from a generalized scorpaenid ances- 

 tor. One of us (WNE) has noted that the gasbladders of the 

 triglids and dactylopterids appear to be similar, with anterior 

 and posterior lobes and very large intrinsic muscles (sec Evans, 

 1973, for information on triglids). In dactylopterids the gas- 



