STEVENS ET AL.: AMMODYTOIDEI 



575 



Table 148. Selected Pigment Characters of Ammodytidae Larvae. = absent, + = present, 



with development, po = posterior, an = anterior. 



increasing with development, i = decreasing 



Species 



Stage 



Body length 



(mm) Jaws 



Snout Brain 



Nape 



Dorsal 

 midline 



Fin- 

 Caudal fold 



Gymnammodyles semisquamaliis prefiexion 



flexion 

 postflexion 



prefiexion 

 postflexion 



flexion 

 postflexion 



G. cicerellus 



Hyperoplus lanceolatus 



H. immaculatus 



Ammodyles tobianus 



A. marinus 

 dubius 

 americanus 



A. hexapterus 



preflexion 



flexion 



postflexion 



preflexion 



flexion 



postflexion 



preflexion 



flexion 



postflexion 



preflexion 



flexion 



postflexion 



4.8 



7.0 



11.8-38.0 



5.5-9.0 

 13.0 

 26.0 



4.0-5.0 



7.5-12.0 



16.0-27.0 



4.5-6.0 



7.5-11.0 



19.0-33.0 



7.0-8.0 

 11.0-13.0 

 16.0-31.0 



7.0 



11.0-25.0 



6.0 



11.0-25.0 0, + 





 



+ 





 

 





 

 





 



OT 





 

 OT 







+ ,0 



+ 





 



+ .0 







+ 



+ 



0, + + 



01 



+ 



+ + 







+ 



+,0 







+ 

 + 





 

 01 



+ 



+1 





 



+ 



+1 



an 

 an 

 an 



+ an 



+ ,0 an 

 + ,0 



an 

 an 



+ 



an 

 an 

 an 





 near tail 



T + 



po 1/3 



+ 



po 1/4 





 near tail 



I 





 po 1/4 



I 





 



01 







+ 

 + 



+ 

 + 







I 







+ 

 + 





 



01 



+ 

 + 

 + 







+ 

 + 



Cameron, 1959 

 Macer, 1967 



Page, 1918 



Einarsson, 1951 

 Macer, 1967 



Macer, 1967 



Einarsson, 1955 

 Macer, 1967 



Einarsson. 1951 

 Macer, 1967 



Kobayashi, 1961c 

 NWAFC, unpubl. 



species, and transformation to juveniles occurs at about 40 mm. 

 The caudal fin is the first to ossify, followed by the pectorals, 

 then the dorsal and anal. The median fin rays form in the pos- 

 terior part of the body, and ossification proceeds forward. Dur- 

 ing larval development the body thickens somewhat, but main- 

 tains its elongate shape. All adult Ammodytidae have protrusible 

 upper jaws, but Gymnammodytes semisquamatus is the only 

 species in which this character is reported in larvae as small as 

 9 mm (Cameron, 1959). Postflexion larvae oi Hyperoplus de- 

 velop vomerine teeth which persist in the adult, while Gym- 

 nammodytes postflexion larvae develop both vomerine and pre- 

 maxillary teeth which disappear at about transformation. During 

 the larval period Gymnammodytes. Hyperoplus. and .Ammo- 

 dyles are pelagic. Juveniles and adults are both pelagic and 

 benthic. 



Pigment. — Pigment can be a useful diagnostic feature among 

 the larvae of Ammodytidae, especially the location and devel- 

 opment of melanophores on the ventral gut margin, the dorsal 

 body margin, and the caudal area, i.e., the tip of the notochord 

 and the edge of hypural elements. These pigment characters are 

 summarized in Table 148. All species have a row of melano- 

 phores dorsally on the gut, beginning at or just posterior to the 

 cleithrum, and a postanal row on the ventral body margin from 

 the anus to the tail. The dorsal gut pigment becomes obscured 

 with growth. Specific variations in pigment patterns can be seen 

 in the 16 mm specimens illustrated in Fig. 304. At this length, 

 dorsal midline pigment forms a complete row in H. lanceolatus, 

 but occurs only on the posterior quarter in .-i. hexapterus and 

 .4. marinus: and ventral gut pigment extends the length of the 

 gut in H. lanceolatus and .4. marinus but is found only on the 

 anterior ventral gut of .4. hexapterus. Pigment patterns of .1. 

 marinus, .4. dubius and .4. americanus are nearly identical (Ma- 

 cer, 1976) although Richards (1982) has noted diflTerences in the 

 ranges of melanophore numbers, especially on the anterior ven- 

 tral gut (stomach) and dorsal midline (supradorsal). Pigment 



appears variously on the head, increasing with age in all species 

 reported. The only reported decrease in pigmentation is on the 

 dorsal and ventral margins of G. cicerellus (Page, 1918). G. 

 semisquamatus has pigment on the ventral finfold margin, the 

 only ammodytid species for which finfold pigment has been 

 noted (Cameron, 1959). 



Aferistics. — Fin ray and vertebral counts for the family Am- 

 modytidae are: Vert 54-78; D 40-69; A 14-36; Pec. 10-16; Pel. 

 0-1,5; and C 15-17 prin., 1 3 branched. In all genera the number 

 of precaudal vertebrae exceeds the number of caudal. Robins 

 and Bohike (1970) report 9 + 8 principal caudal rays for Em- 

 bolichthys sarissa. but all other ammodytid species, including 

 E. mitsukurit, have 8 + 7. Embolichthys is the only genus with 

 pelvic fins, which are thoracic. The caudal fin is the first to form, 

 followed by the pectorals, dorsal, and anal. Posterior rays of the 

 median fins form first and development proceeds forward. Fin 

 formation is completed by 30 to 40 mm body length. 



Relationships 



Although early life history data of the suborder Ammody- 

 toidei do little to clarify its phylogenetic position, larval pigment 

 patterns and myomere- vertebrae counts are useful in separating 

 sympatric species (Macer, 1967). General characters, such as 

 the well developed state of newly hatched larvae and the se- 

 quences of fin development, are shared with other perciform 

 derivatives and relatives, but essentially the problem of the 

 systematic position of ammodytids is not yet resolved. 



(A. CM.) National Marine Fisheries Service, Northwest 

 AND Alaska Fisheries Center, 2725 Montlake Boule- 

 vard East, Seattle, Washington 98112; (E.G.S.) Na- 

 tional Marine Fisheries Service, Southwest Fisheries 

 Center, PO Box 27 1 . La Jolla, California 92038; (W.W.) 

 Marine Ecological Consultants, 531 Encinitas Boule- 

 vard, Slute 1 10, Encinitas. California 92024. 



