The grenadier anchovy, Coilia, enters the LRT alongside the anchovy, Engraulis, but far from the herring, Clupea

Updated June 2, 2023 with a new nesting
of Coilia (Figs 1, 2) and Engraulis (Figs 3, 4) in the Palaeonisciformes. The anterior extension of the nasals (pink) is a trait this clade shares. Click here for more data on this shift based on adding taxa and greater undertanding of the facial bones.

Figure 1. The Japanese grenadier anchovy, Coilia nasus.

Everyone agrees
the grenadier anchovy (Coilia, Figs. 1, 2) and the anchovy (Engraulis, Figs. 3, 4) are closely related taxa. But when you add taxa, anchovies nest apart from herring, their traditional sisters in the clade Clupeiiformes. Now these two taxa nest within the Palaeonisciformes, a clade just prior to the lobefin-ray fin split in the Osteichthyes in the large reptile tree (LRT, 2264 taxa).

Figure 2. Skull of Coilia nasus, the Japanese grenadier. Note the outrageous maxilla.

Coilia nasus
(Gray 1830; up to 41cm) is the Japanese grenadier anchovy, commonly found at 50m depths, but migrates to freshwater to spawn. Others live in freshwater lakes continually. Note the insanely long maxilla extending beyond the back of the skull. There is nothing else like it in the Vertebrata. The incurrent nares appear to open ventrally, below the nasals.

Figure 2. Engraulis the anchovy.
Figure 3. Engraulis mordax, the anchovy.
Figure 4. Engraulis skull greatly resembles that of Clupeopsis.

Compare
Coilia to Engraulis, the anchovy (Figs. 2, 3), a more plesiomorphic taxon. Everything is a little weirder in Coilia. Note the breakup of the postorbital (amber) in Coilia. This is a neotonous trait. Juveniles don’t have fused facial bones.

References
Gray JE 1830. : Illustrations of Indian Zoology 1, (pl. 85).

wiki/Coilia
wiki/European_anchovy
wiki/Coilia
wiki/Engraulinae

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