582 



ONTOGENY AND SYSTEMATICS OF FISHES-AHLSTROM SYMPOSIUM 



A definitive phylogenetic reconstruction of zoarceoid rela- 

 tionships is not presently possible without a more thorough 

 knowledge of the anatomy of other fishes that have been tra- 

 ditionally allied with them. Preliminary phylogenetic inferences 

 were made by Anderson (1984), who also discussed relation- 

 ships among zoarcid genera. It should be noted, however, that 

 a search for more characters is still in progress. Makushok (1958) 

 and Springer (1968) suggested zoarceoid, or "northern blen- 

 nioid" relationships were not close to the "tropical blennioids," 

 a fact that my own research supports. However, for the con- 

 venience of the reader, information on the early life history 

 stages of zoarceoids, excluding Zoarcidae, is given by Matarese 

 et al. (this volume) under Blennioidea, following Nelson ( 1 976). 



Since there is a dearth of knowledge on early stages and since 

 the youngest specimens known of any zoarcid so closely resem- 

 ble adults, no early life history characters have helped in elu- 

 cidating systematic relationships within Zoarcidae, or the zoar- 



cids to their allies. All the zoarceoids are characterized by 

 precocious early stages (see Matarese et al., this volume), but 

 the utility of these forms in phylogeny remains untested. Within 

 Zoarcidae, it is interesting to note that the development of ce- 

 phalic lateralis pores in the primitive Gymnelus viridis. Melan- 

 ostigma pammelas. and Macrozoarces americanus takes place 

 over a much longer growth period (up to 50-60 mm) than in 

 the more derived Bothrocara (Okiyama, 1982a) or in youngest 

 stages I studied of Lycenchelys (32 mm), Lycodapus (20 mm), 

 or Lycodes (38 mm). The value of this information awaits more 

 complete data on early life history stages of all zoarcids. 



Virginia Institute of Marine Science, College of William 

 AND Mary, Gloucester Point, Virginia 23062. Present 

 Address: Department of Ichthyology, California 

 Academy of Sciences, Golden Gate Park, San 

 Francisco, California 941 18. 



Gobioidei: Development 



D. RUPLE 



GOBIOIDS are one of the most speciose groups of fishes, 

 comprised of approximately 2,000 species or ten percent 

 of the total number of teleosts in the world (Cohen, pers. comm.). 

 Various workers have recognized from two to seven major fam- 

 ilies of gobioids, based on adult characters. For present purposes 

 I will recognize seven families' Eleotridae, Gobiidae, Rhyacich- 

 thyidae, Kraemeriidae, Gobioididae, Trypauchenidae, and Mi- 

 crodesmidae after Nelson (1976). 



Development 



Larvae are known for less than 5% of gobioid species. Eggs 

 and larvae are best known from Japanese waters (e.g., Dotsu. 

 1954, 1957, 1958, 1979; Dotsu and Fujita, 1959; Dotsu and 

 Mito, 1955, 1963; Dotsu and Shiogaki, 1971; Kobayashi et al., 

 1973; Shiogaki and Dotsu, 1971e, 1972c). in the northeastern 

 Atlantic and Mediterranean Sea area (e.g., Petersen, 1917, 1919; 

 Fage, 1918; Lebour, 1919; Sparta, 1934; plus summaries in 

 Padoa, 1956f; and Russell, 1976), and less so in American waters 

 (e.g., Hildebrand and Cable, 1938; Perlmutter, 1939; Pearson, 

 1 94 1 ; and Ruple, in prep.). Most of these descriptive works deal 

 with the gobiids, although larvae are known for representatives 

 of all families except Rhyacichthyidae and Kraemeriidae. 



Larvae of gobioids are fairly distinctive from other teleosts, 

 but considerable variation does occur within the suborder. The 

 diversity of characters found in eggs and larvae will be discussed 

 in the following section. This information was compiled from 

 published literature and the examination of gobioid larvae. 



' Hoese (this volume) includes Gobioididae and Trypauchenidae in 

 the Gobiidae subfamily Amblyopinae and recognizes Xenisthmidae as 

 a distinct family. Eleotridae is changed to Eleotrididae. 



Eggs 



Eggs are known for eleotrids, gobiids, gobioidids, and micro- 

 desmids (Table 151). Eggs of eleotrids and gobiids are generally 

 ellipsoid and adhesive, many of which have filamentous strands. 

 Eggs range in size from as small as 0.40 x 0.32 mm in Eleotris 

 avycep/jfl/a (Eleotridae; Dotsu and Fujita, 1959) and 0.45 x 0.20 

 mm in Evorthodus lyricus (Gobiidae; Foster and Fuiman, MS 

 in prep.) to 3.8 x 1.3 mm in Pcrcottus glehni (Gobxiiisie:; Kry- 

 zhanovsky et al., 1951) and 5.5 x 0.9 mm in Acanthogobius 

 Jlavimanus (Gobiidae; Dotsu and Mito, 1955). Taenioides ruh- 

 icundus (Gobioididae) eggs are demersal, adhesive and measure 

 approximately 1.3 x 0.70 mm (Dotsu, 1957) whi\e Gunnellich- 

 ihys (Microdesmidae) eggs are spherical (Smith, 1958a). 



Known gobioid eggs usually contain numerous small oil drop- 

 lets within the yolk. Newly hatched larvae range from 1.7 mm 

 in Aslerropteryx semipunctatus (Eleotridae; Dotsu and Mito, 

 1963) to 7.0 mm in Chaenogobius castanea (Gobiidae; Dotsu, 

 1954). 



Larvae 



Gross morphology. — Body shape of gobioids is generally slightly 

 elongate and slender, with body depth usually nearly uniform 

 rather than sharply tapering (Figs. 309-311). Gobioidid and 

 microdesmid larvae are moderately elongate and slender (Fig. 

 311), while most eleotrids and trypauchenids are only slightly 

 elongate and slender (Fig. 311). Microdesmids have the most 

 elongate body shape of any known gobioid larvae. Body form 

 within the gobiids exhibits the greatest variety, ranging from 

 fairly short and stout (Gobiidae Larva 1 , Fig. 309) to moderately 

 elongate and slender Luciogobius elongatus (Fig. 309). These 

 characteristic body shapes are usually retained from the larval 

 through adult stages. 



