Blennioidea: Development and Relationships 

 A. C. Matarese, W. Watson and E. G. Stevens 



THE Blennioidea is composed of 16 families with about 182 

 genera and 759 species (Table 143). The families discussed 

 here are those included in the infraorder Blennioidea by Nelson 

 (1976), as amended by the current literature. For convenience 

 we divide the infraorder into a tropical and a northern group. 

 The tropical group is similar to Gosline's ( 1 968) superfamily 

 Blennioidae except for the following: 1) Ophiclinidae and Per- 

 onedysidae are synonymized with the Clinidae (George and 

 Springer, 1980); 2) Dactyloscopidae is included (George and 

 Springer, 1980); and 3) Congrogadidae is excluded (Winterbot- 

 tom, 1982'). The northern group is similar to Gosline's (1968) 

 superfamily Zoarceoidae except that we include the Bathymas- 

 teridae (Anderson, 1984). The Zoarcidae is treated separately 

 (Anderson, this volume). 



The majority of species (80%) belong to four tropical families: 

 Tripterygiidae, Clinidae, Labrisomidae, and Blenniidae. Of the 

 northern forms, only the family Stichaeidae represents a sig- 

 nificant percentage (8%) of the species. Tropical Blennioidea 

 inhabit primarily the Indo-West Pacific south to Australia, while 

 northern fishes inhabit the North Atlantic and North Pacific 

 (Table 143). Occasionally, representatives of mainly tropical 

 families occur in boreal waters (e.g., Clinidae and Blenniidae), 

 and members of northern families may occur in the subtropics. 

 Some dactyloscopids inhabit fresh water. Four families are 

 monotypic and three of these are endemic to the northeast Pa- 

 cific. 



As a group most of the 16 families in Blennioidea are not 

 well understood, probably due to their lack of commercial im- 

 portance, small size and cryptic habits. In general, the tropical 

 and more speciose families (e.g., Blenniidae) are better known 

 than the northern families. Monotypic families are quite poorly 

 known. Although sparse and incomplete, some early life history 

 information is available for II of the 16 families (Table 143). 

 In most cases, however, the data on few species may not be 

 representative of the family. Among the families in the infraor- 

 der, the Blenniidae has the greatest number of species (22) de- 

 scribed; but with about 319 species in the family, this amounts 

 to fewer than 10%. Morphology, pigment, and meristics of lar- 

 vae in the infraorder are diverse (Figs. 302, 303). 



Development 



Eggs 



Fishes in this infraorder spawn demersal eggs (Table 144), 



except for some clinids. Clinids of the tribe Ophiclinini are 



ovoviviparous (George and Springer, 1980), while those of the 



tribe Clinini are viviparous (Penrith, 1969; Hoese, 1976). 



Most blennioid eggs are spherical to somewhat flattened, pos- 

 sess one to several oil droplets, are attached to one another (and 

 often to a substrate) by filaments or other adhesions, and have 



' Winterbottom, R. 1982. The perciform fish family Congrogadidae— 

 biogeography and evidence for monophyly. Amer. Soc. Ich. Herp., oral 

 paper, 62nd annual meeting. 



a smooth unsculptured chorion. Sizes range from among the 

 smallest offish eggs (Blenniidae, 0.50 mm) to among the largest 

 (Anarhichadidae, 8.0 mm). Incubation periods range from 6 to 

 70 days. Eggs are unknown for four families: Xenocephalidae, 

 Ptilichthyidae, Zaproridae, and Scytalinidae. 



Parental care is common among most families; e.g., in sti- 

 chaeids. males or females may guard egg masses (Shiogaki and 

 Dotsu, 1972a; Shiogaki, 1981, 1982). In an extreme example 

 of parental care, male dactyloscopids incubate eggs in ball-like 

 clusters carried beneath the pectoral fins (Dawson, 1982). 



Larvae 



Morphological characters— ^Xenmoidea larvae hatch at sizes 

 ranging from as small as 2.0 mm (Blenniidae) to as large as 1 7.0 

 mm (Anarhichadidae) (Table 145). Larvae of the northernmost 

 families hatch at more than twice the size of larvae of the more 

 tropical families (i.e., averaging ca. 1 1.5 mm versus ca. 4.5 mm). 

 Size at which notochord flexion is complete is also variable, but 

 tropical larvae are usually fully flexed by ca. 10.0 mm whereas 

 northern larvae do not complete flexion until ca. 20.0 mm. At 

 least three families have larvae with an extended pelagic exis- 

 tence: Blenniidae, Cryptacanthodidae, and Zaproridae. Mem- 

 bers of the blenniid tribe, Salariini, have the only well-docu- 

 mented, prejuvenile pelagic stage (Miller et al.. 1979; Leis and 

 Rennis, 1983). This has been termed the "ophiblennius"" stage 

 and usually occurs between 4.6 and 26.0 mm (Fig. 302). At least 

 two families, Cryptacanthodidae (Shiogaki, 1982) and Zapror- 

 idae (Haryu and Nishiyama, 1981), have heavily pigmented 

 larvae and prejuveniles that are extensively collected in surface 

 nets suggesting an extended pelagic existence (Fig. 303C, G). 

 Most blennioid larvae do not undergo a marked metamorphosis. 

 Transformation is usually complete in tropical forms by 26.0 

 mm, but may begin as early as 10.0 mm in some families (Trip- 

 terygiidae and Blenniidae). Larvae in the more northern families 

 transform at a slightly larger size, ca. 30.0-40.0 mm, although 

 Ptilichthys transforms at ca. 1 14.0 mm. 



Among the tropical families, larval Tripterygiidae, Clinidae, 

 and Labrisomidae share many similar morphological features. 

 They are moderately elongate, have a preanal length about 50% 

 BL (slightly less in labrisomids), possess a large swimbladder, 

 and usually lack preopercular spines (Figs. 302A, B, C, D). 

 Heads are small, sometimes rounded, with a short snout. Mouths 

 extend just beyond the anterior eye margin. In tripterygiid and 

 clinid larvae, the gut is initially straight but coils during flexion. 



The blenniids include many larval forms with diverse mor- 

 phological features. According to Leis and Rennis (1983), how- 

 ever, larvae are more similar within tribes than between tribes. 

 Most species are moderately elongate (Nemophini includes both 

 slender and robust forms), becoming either more slender (Nem- 

 ophini) or more robust (Salariini) with development. Heads are 

 short, rounded, and broad becoming more elongate with de- 

 velopment (except Salariini larvae in which the snout elongates 

 early in the preflexion stage). The gut is short to moderate (usu- 

 ally < 50% BL), and eventually coiled if not so initially. L-arval 

 preopercular spination may be elaborate: spines can be numer- 



565 



