FRITZSCHE: GASTEROSTEIFORMES 



405 



thoids (sic) suggest one or two origins in the percopsiform — 

 beryciform area." The mixture of advanced and primitive char- 

 acters shown by gasterosteiforms suggested to Banister (1967) 

 evolution "from a primitive myctophoid type offish . . . towards 

 an acanthopterygian grade." McAllister (1968) suggested "the 

 Gasterosteiformes are derivable from the Perciformes" and ". . . 

 the Syngnathiformes from the subperciforms, such as Beryci- 

 formes and Zeiformes." In fact he suggests that Antigonia or 

 Capromimus would appear to be close to the ancestors of the 

 Syngnathiformes. None of these authors presented evidence for 

 support of their ideas. Examination of the description o^ Anti- 

 gonia larvae by Nakahara ( 1 962) shows that this fish bears little 

 resemblance to the early stages of described gasterosteiforms. 

 Larval Antigonia are characterized by well-developed, serrated 



preopercular and cranial spines. These spines are never seen in 

 gasterosteiform larvae. However, the description of the larvae 

 of Capros aper (Russell, 1976) indicates that the most charac- 

 teristic feature of them is the occurrence of small spines all over 

 the body surface. Additionally the larvae of C. aper are darkly 

 pigmented. These two characteristics are also found in some 

 gasterosteiform larvae, e.g., Macrorhamphosus. It is therefore 

 tempting to use these characters in support of McAllister's hy- 

 pothesis, however we will have to wait for further information 

 on both gasterosteiforms and zeiforms before we can support 

 or refute this hypothesis. 



Department of Fisheries, Humboldt State University, Ar- 

 CATA, California 95521. 



Scorpaeniformes: Development 

 B. B. Washington, H. G. Moser, W. A. Laroche and W. J. Richards 



THE Scorpaeniformes are the fourth largest order of fishes 

 encompassing about 20 families (depending on classifi- 

 cation used), 250 genera and over 1 ,000 species. Representatives 

 of the order are widely distributed from tropical to arctic and 

 antarctic waters. Most scorpaeniforms are benthic or epibenthic 

 with representatives ranging from freshwater to the deep ocean. 



The morphologically diverse "mail-cheeked fishes" are named 

 for the bony suborbital stay which extends posteriorly from the 

 third infraorbital to the preopercle. The suborbital stay is the 

 only known character that defines the order; however, some 

 workers have suggested that the stay evolved independently in 

 several lineages and may not indicate monophyly (Matsubara, 

 1943; Quast, 1965; Poss, 1975). The classification of the scor- 

 paeniforms is controversial, not only in terms of monophyly 

 but also at the subordinal and familial levels. Discussion of the 

 taxonomic status and current hypotheses of relationships is pre- 

 sented in Scorpaeniformes: Relationships (this volume). 



Modes of reproduction vary widely within the scorpaeni- 

 forms. Many families spawn individual pelagic eggs (Anoplo- 

 pomatidae, Congiopodidae, Hoplichthyidae and Triglidae), while 

 others spawn demersal clusters of adhesive eggs ( Agonidae. Cot- 

 tidae, Cyclopteridae and Hexagrammidae). Where known, most 

 scorpaenids produce pelagic egg masses enclosed in a gelatinous 

 matrix. Notable exceptions include the scorpaenid genus Se- 

 bastes and the comephorids of Lake Baikal which give birth to 

 live young. 



Larvae of only about 20% of scorpaeniform genera and ap- 

 proximately 10% of the species are known. Because of the wide 

 diversity of form, we are not able to characterize a typical scor- 

 paeniform larva. Early life stages of many scorpaeniforms are 

 characterized by strong head spination as depicted in the gen- 

 eralized scorpaenid larva Sebastes (Fig. 220). However, the 

 expression of head spination is variable within the order with 

 elaborations and losses in many groups. 



For the purposes of this paper, we consider the Scorpaeni- 

 formes to be monophyletic and utilize the broad suborders Scor- 



paenoidei and Cottoidei as a framework for presentation and 

 discussion. Because of the order's morphological diversity and 

 the lack of an agreed upon classification, discussion of larval 

 taxonomy is focused upon each family. The scorpaeniform fam- 

 ily Cyclopteridae is presented in the subsequent article in this 

 volume. 



SCORPAENOIDEI 



Eggs 



Eggs are known for seven of the scorpaenoid families recog- 

 nized in Washington et al. (this volume), however, they are 

 known only for a few species (Table 107). Most scorpaenoid 

 families are oviparous and spawn pelagic eggs; however, repro- 

 ductive modes are varied in the Scorpaenidae. In the scorpaenid 

 subfamilies Scorpaeninae, Pteroinae, and Sebastolobinae the 

 eggs are extruded in bilobed gelatinous egg masses which float 

 at the surface. The eggs are slightly elliptical and have homo- 

 geneous yolk, a narrow perivitelline space, and a smooth cho- 

 rion. A single oil globule is present in Pterois (0.16-0.17 mm) 

 and Sebastolobus (0. 18-0.20 mm); Scorpaena lacks an oil glob- 

 ule. In the choridactyline genus Inimicus, eggs are extruded 

 singly, are spherical, and lack an oil globule (Table 107). Mem- 

 bers of the scorpaenid subfamily Sebastinae are viviparous and 

 give birth to large broods of young which are comparable in 

 stage of development to first-feeding larvae of oviparous scor- 

 paenids. The eggs are retained in the lumen of the ovary after 

 ovulation, range between 0.75 and 1.9 mm, have homogeneous 

 yolk, a narrow perivitelline space, smooth chorion, and one to 

 many oil globules. For the other families for which eggs are 

 known, the eggs are pelagic with none to multiple oil globules 

 (Table 107). 



Larvae 



At least one larval stage is known for 64 of the more than 

 600 species of scorpaenoids and for 20 of the 100+ genera. 

 Major reviews of larval scorpaenoids include Sparta (1956b) 



