WASHINGTON ET AL.: SCORPAENIFORMES 



439 



and gobioids) should not be excluded from consideration of 

 relationships to some scorpaeniform groups. 



In summary, the limits of the order, suborders, families and 

 distribution of families in the suborders are the subject of con- 

 siderable disagreement among current workers. These problems 

 will not be resolved without a worldwide revision of the order. 

 At this point, we assume that the order is monophyletic. For 

 the purposes of summarizing information on this order, we treat 

 two broad suborders: the Scorpaenoidei and the Cottoidei. We 

 consider these groups as a convenient way to discuss disagree- 

 ments in classification of specific groups and hypotheses of re- 

 lationships; we do not propose that they are monophyletic groups. 



Suborder Scorpaenoidei 



For this paper, we recognize the Scorpaenoidei to include the 

 following families: Scorpaenidae (broad sense of Matsubara, 

 1943), Triglidae, Peristediidae, Bembridae, Platycephalidae, 

 Hoplichthyidae, and Dactylopteridae. Some of these families 

 have been assigned to separate suborders or superfamilies and 

 the dactylopterids have often been placed in a separate order 

 (Quasi, 1965; Nelson, 1976; Lauder and Liem, 1983). 



Meristic features and approximate number of species for in- 

 cluded groups are provided in Table 111. Data have been drawn 

 from many sources and may not be complete for some genera 

 or may omit extremes found in abnormal individuals. 



Matsubara's work (1943) is the most thorough study of scor- 

 paenoids to date. His hypothesis of relationships (Figure 240) 

 is based on a wide variety of characters including those of the 

 infraorbital bones, suspensorium, hyoid apparatus, cranium, 

 pectoral girdle and gasbladder. Matsubara included 14 subfam- 

 ilies in his family Scorpaenidae. He recognized three large ge- 

 neric groups or lineages within the scorpaenoids which he la- 

 beled: Sebasles-slem, Scorpaena-siem and Cocotropus-stem. His 

 Sehasles-stem contains two subfamilies, the Sebastinae and 

 Neosebastinae which were viewed as the most primitive or "gen- 

 eralized" of the scorpaenoids. His second group, the Scorpaena- 

 stem, includes five subfamilies: Scorpaeninae, Pteroinae, Setar- 

 chinae, Sebaslolobinae, and Plectrogeninae. The third group, 

 the Cocotropus-ilem, includes six subfamilies: Apistinae, Con- 

 giopinae, Aploactinae, Minoinae. Pelorinae, and Erosinae. The 

 latter two groups were considered "specialized" or derived rel- 

 ative to the Sebastes-%\em. Other worJcers (Greenwood et al., 

 1966; Nelson, 1976; Poss and Eschmeyer, 1978) have departed 

 from Matsubara's classification of the Scorpaenidae by elevating 

 some subfamilies of Matsubara to family status. In addition, 

 the Congiopodidae [but not Matsubara's Congiopinae (sic)] has 

 been recognized as a separate family in a monotypic suborder 

 by Greenwood et al. (1966) and Nelson (1976) and as a super- 

 family by Quast (1965). Other scorpaenoid groups not treated 

 by Matsubara (1943) have been given separate status within the 

 Scorpaenoidei by the aforementioned workers, and include the 

 Caracanthidae and Pataecidae. In his later work on fish hier- 

 archy, Matsubara (1955) recognized three families of scorpae- 

 nids which basically correspond to his earlier three "stem" groups. 



We basically follow the phylogenetic hypotheses of Matsubara 

 (1943, 1955) in presenting general trends in relationships within 

 the suborder. The following discussion highlights groups where 

 problems or disagreements about relationships are persistent. 

 A phylogenetic approach based on information presented here 

 would result in family and subfamily lines being interpreted 

 quite differently. However, we believe presentation of a new 



Fig. 240. Schematic representation of scorpaeniform relationships 

 from Matsubara ( 1 943). 



classification is premature; a thorough study of the scorpaenoids 

 is required on a worldwide basis. 



The Sebastinae is currently considered to be the most prim- 

 itive or generalized group of scorpaenoids because of the in- 

 complete suborbital stay in Sebastes, weak head spination, and 

 general body plan similar to the percoids (Matsubara, 1943 and 

 others). Matsubara (1943) proposed that Sebastes was the most 

 generalized genus within the subfamily with a transition series 

 to Helicolenus. Eschmeyer and Hureau (1971) and Barsukov 

 (1973) believed that Matsubara's transition series is reversed 

 with Helicolenus the most generalized genus and Sebastes being 

 a relatively derived form. 



The subfamily Scorpaeninae with its 1 50 genera is considered 

 a "catch-basket" subfamily, and there is no certainty that it is 

 monophyletic. 



Matsubara (1943) noted that the Setarchinae lack a basi- 

 sphenoid as do cottoids and that the second and third actinosts 

 intervened between the hypercoracoid and hypocoracoid. He 

 concluded from these observations that the Setarchinae and 

 cottoids shared a common ancestor. However, Eschmeyer and 

 Collette (1966) disagree. In their review of the Setarchinae, a 

 small basisphenoid, connected only by cartilage, was found in 

 cleared and stained specimens; they stated that Matsubara's 

 conclusion was untenable. 



Matsubara (1943) suggested that the genus Sehastolobus was 

 closely related to the genus Plectrogenium (subfamily Plectro- 

 geninae) because of their shared lack of gasbladders, notched 

 pectoral fins and prominent rows of spines along the sides of 

 their head. He further noted (1943:160): thai "Plectrogenium 



