A Hypomaxillary Bone in Harengula (Pisces: Clupeidae) 
Frederick H. Berry ] 
The herring genus Harengula Valenciennes 
(as herein restricted) contains five bilateral 
pairs of bones in the upper jaw (Fig. 1 A). 
Most other clupeid fishes contain three or four 
such pairs of bones: premaxillary, maxillary, 
and one or two supramaxillaries. The extra pair 
of bones in Harengula is here termed the hypo- 
maxillary. The hypomaxillary also occurs in the 
clupeid genera Pliosteostoma Norman and Pel- 
lona Valenciennes, and its presence has been 
used to distinguish these two genera from other 
genera. The presence of the hypomaxillary in 
Harengula and its usefulness as a taxonomic 
character in separating Harengula from other 
closely related genera previously has been over- 
looked. 
The hypomax illary previously has been 
termed "an ossified ligament,” "a narrow, toothed 
bone,” and "a bone, bearing teeth lying in the 
membranous section between the distal end of 
the premaxillary and the middle of the maxil- 
lary.” It is desirable to have a specific name for 
this unique pair of bones, and hypomaxillary is 
a logical choice. Other similar terms that have 
restricted meaning or are synonyms of other 
bones are intermaxillary, inframaxillary, bimax- 
illary, submaxillary, and surmaxillary. 
Four closely related genera are involved in 
the evaluation of this character: Harengula 
1 U.S. Bureau of Commercial Fisheries, Biological 
Laboratory, Brunswick, Georgia. Manuscript received 
Valenciennes 1847, Lile Jordan and Evermann 
1896, Sardinella Valenciennes 1847, and one 
whose designation currently is uncertain, but 
here is referred to as Clupalosa Bleeker 1851. 
( The distinguishing characteristics are listed 
below. ) 
The position of the hypomaxillary in Haren- 
gula is shown in Figure 1 A. Its posterior end 
overlaps the maxillary laterally. The hypomaxil- 
lary, premaxillary, and maxillary all bear a single 
row of small pointed teeth. The connective 
tissue in the space between the hypomaxillary 
may also bear teeth. The hypomaxillary was 
ossified in the smallest specimen examined (16 
mm S.L.). The presence of the hypomaxillary 
and the characteristic elongation of the posterior 
supramaxillary (Fig. 2 A) in Harengula have 
been verified in the following species: 
H. clupeola (Cuvier 1829), St. Lucia, British 
West Indies, SU35458, and syntype of H. 
latulus Valenciennes, SU32769 
H. burner alts (Cuvier 1829), Jamaica, SU5041 
H. pensacolae Goode and Bean 1880, Sanibel 
Island, Florida, SU36092, and Santos, Brazil, 
SU36065 (as S. majorina Storey 1938) 
H. peruana Fowler and Bean 1924, many 
specimens from Peru to Costa Rica in 
several institutions 
H. thrissina Jordan and Gilbert 1882, many 
specimens from Pacific Mexico in several 
August 1, 1963. 
institutions 
HYPOMAXILLARY 
SHAPE OF 
POSTERIOR 
SUPRAMAXILLARY 
BILOBED FOLD 
ON CLEITHRUM 
Harengula 
present 
elongated, ventral 
constriction anterior 
present 
Lile 
absent 
elongated, ventral 
constriction anterior 
absent 
Sardinella 
absent 
rounded, ventral constriction 
vertical to dorsal constriction 
present 
Clupalosa 
absent 
elongated, ventral 
constriction anterior 
present 
373 
