McGOWAN AND BERRY: CLUPEIFORMES 



123 



(Wongratana, 1 983) and provided keys to all Indo-Pacific species 

 (Wongratana, 1980). However no direct comparison between 

 his classification and that of Nelson (1967, 1970, 1973) is pos- 

 sible because he only examined Indo-Pacific material while Nel- 

 son included West African and New World material. 



Evidence from eggs and larvae 



There are two major problems with using characters of eggs 

 and larvae to criticize classifications based on adult characters. 

 First, the planktonic stages of fishes are exposed to different 

 selective pressures than the adults so they may show patterns 

 of specializations for planktonic life which are not congruent 

 with the distribution of adult character states. Second, relatively 

 few genera of clupeiform fishes have had the eggs or larvae 

 described for even one species in the genus. The first problem 

 must be dealt with the same as any character complex in a group 

 with more than one character complex. More knowledge of the 

 ecology of the larvae in the sea would indentify species with 

 different funtional requirements for their larvae. The second 

 problem may be resolved by using the available evidence in a 

 parsimonious fashion. 



Eggs and young larvae are similar within genera. Seven species 

 of Sardine/la (Table 25) all have moderately sized clupeid-type 

 eggs with a wide perivitelline space and a single oil globule. The 

 egg described by Takita (1966) and Chang et al. (1981) as that 

 oi Harengida ziinasi is similar. Wongratana ( 1 980) places zunasi 

 in Sardinclla. 



Within subfamilies there is little apparent consistency in egg 

 morphology among genera. Etruineus has no oil droplet but 

 Dussumieria does. Brevoortia has eggs 1.3 mm or larger with a 

 single oil globule; HHsa kelee has 1.00-1.07 mm eggs with sev- 

 eral small oil droplets. Clupea has demersal adhesive eggs while 

 Sprattus has pelagic eggs with a small perivitelline space. The 

 Indo-Pacific pristigasterin species of Ilisha have large eggs with 

 adhesive coatings and a single large oil globule but Opislhopterus 

 tardoore and the eastern Pacific O. dovii have small eggs with 

 small perivitelline spaces and no oil droplets. 



The functional significance of egg characters is unknown. Sep- 

 arate lineages within the group which have radiated into several 

 habitats could show parallel adaptations such as oil droplets for 

 buoyancy or nutrition, adhesive coating for retention nearshore 

 or demersally. and egg size as a trade-off between broadcasting 

 and parental investment. Alternatively, different types of eggs 

 within taxonomic categories could also support splitting the 

 category. The anchovy genus Stolephorus contains species with 

 eggs which range from oval with no oil globule to varying degrees 

 of eccentricity with an oil droplet, to unusually shaped eggs with 

 knobs on one end (Delsman, 1931). Nelson (1983) separated 

 Stolephorus into two groups, a Stolephorus group with 1 3 species 

 and an Encrasicholina (new usage) group of 5 species which he 

 considered more closely related to New World anchovies than 

 to the 1 3 Stolephorus species. The three Encrasicholina species 

 whose eggs are known have an oval egg without a knob. One 

 of the three, E. hetcrolobus. was reported by Delsman (1931) 

 to have a small oil droplet and to be relatively more abundant 

 near shore than Stolephorus zolingeri. The other two, E. pur- 

 purcus and E. punctifer (^buccanceri, Strasburg, 1960; =zolin- 

 geri. Delsman, 1931), occur in Hawaii and neither has an egg 

 with an oil droplet. New World anchovies don't have eggs with 

 knobs or oil droplets; therefore, the evidence from eggs supports 



Nelson's revision and in addition provides some basis for zoo- 

 geographic speculation. 



Whether the pristigasterins should be given equal rank with 

 the clupeids and engraulids cannot be answered with the avail- 

 able ontogenetic information. There are two very different egg 

 types in the group, small with small perivitelline space and large 

 with gelatinous coating, both of which could be considered spe- 

 cializations. Etrumeus. Jenkmsia. Spratelloides, Clupea. Sprat- 

 tus, and Potamalosa were linked based on a foramen in the 

 fourth epibranchial (Nelson, 1970). Eggs of Spratelloides and 

 Clupea are both demersal and adhesive. The planktonic eggs of 

 Etrumeus and Sprattus both have narrow perivitelline spaces 

 and lack oil globules. Eggs of Potamalosa and Jenkinsia are 

 unknown. Jenkinsia is related to Spratelloides and has demersal 

 larvae (Powles, 1977) so it may have demersal eggs. The de- 

 velopmental osteology of these genera could be studied to de- 

 termine if the shared foramen is phylogenetically homologous. 

 The egg of Anodontostoma, Dorosominae, is similar to eggs of 

 the Alosinae in that it has multiple small oil droplets. Otherwise 

 both the Alosinae and Dorosomatinae contain species with de- 

 mersal adhesive eggs and species with buoyant planktonic eggs. 



Other suggestions of Nelson (1970) that Sardinclla. Opistho- 

 nema. Harengula. and Herklotsichthys should be placed with 

 the Dorosomatinae and Sardina and Sardinops with the Alo- 

 sinae and then that the Alosinae and Dorosomatinae should be 

 combined leaving just Clupeinae and Dorosomatinae cannot be 

 critically evaluated with existing ontogenetic data. These hy- 

 potheses could be tested by comparing the osteological devel- 

 opment of the characters used by Nelson, augmented by other 

 early life history characters. 



Relationships of the Clupeiformes 

 Greenwood et al., (1966) placed the Clupeomorpha and Elo- 

 pomorpha together in their Division One but gave serious con- 

 sideration to the possibility that the Clupeomorpha should be 

 recognized as a separate division. Using information on the gut 

 and lower jaw. Nelson (1973) proposed that the Clupeomorpha 

 were distinct from the Elopomorpha but perhaps related to the 

 non-osteoglossomorph teleosts. Gosline (1980) concluded that 

 the clupeiform fishes should be grouped with the elopiform, the 

 salmoniform, gonorynchiform, and ostariophysine fishes; sep- 

 arated on one side from the osteoglossiform fishes and from the 

 iniomous— acanthopterygian teleosts on the other. His conclu- 

 sions were based on five morphological character complexes: 

 the caudal skeleton, the swim bladder-ear connection, the post- 

 cleithrum, the structures associated with pectoral fin movement, 

 and the various types of premaxillary movements and jaw pro- 

 trusion (Gosline, 1980). 



Gosline (1980) considered the elopomorphs to be an early 

 offshoot from a basal lower teleostean group. He considered the 

 gonorynchiforms and ostariophysines to be more closely related 

 to each other than to the clupeiforms. A clupeiform— osteo- 

 glossiform link has also been mentioned (Greenwood, 1973). J. 

 S. Nelson (1976), who put the superorders Clupeomorpha (Clu- 

 peiformes) and Elopormorpha (Elopiformes, Albuliformes, An- 

 guilliformes) into Division Taeniopaedia, slated succinctly that 

 "the relation of superorders recognized here is poorly known 

 and they are essentially "loose ends." " Lauder and Liem (1983) 

 provisionally follow Nelson (1970) for most groups within the 

 Clupeomorpha but represent the interrelationships of clupeoid 

 lineages as an unresolved polychotomy. Lauder and Liem (1983) 



