Updated October 29, 2022
with a new DGS tracing of Ozarcus now nesting at the base of all sharks. Much of what was written below were valid earlier, with fewer taxa and earlier interpretations, now rendered invalid.
Following the advent of transverse toothless jaws
in Silurian Loganiella and its extant sisters, Rhincodon (the whale shark) and Manta, the Large Reptile Tree (subset Fig. 1) splits taxa with newly acquired anteroposteriorly deeper jaws + marginal teeth in two during the first great dichotomy. At this point the LRT splits elasmobranchii (sharks + ratfish in blue Fig. 1) from stem bony fish like Ozarcus (Figs. 2, 3).
In phase two,
The clade of stem bony fish (that had no bone) quickly and ultimately evolved into genuine bony fish like Amia (the bowfin), at the top of the orange column (Fig. 1) and Trachinocephalus (the lizardfish), at the top of the yellow cladogram (Fig. 1).
Members of this clade and those that directly precede it
are step-wise ancestors to tetrapods, mammals and humans.
The basalmost shark in the LRT
is now Ozarcus here (Figs. 2, 3) shown for the first time with bones colored with tetrapod homologs for scoring. It had large eyes, a deep jawline and the tiniest marginal teeth. Such teeth were a new invention and this is how they started, not much bigger than dermal denticles. It was so close to sharks, it was originally described as “a Palaeozoic shark with osteichthyan-like branchial arches.”
With its giant eyes and short rostrum
Ozarcus also looked like a spiny shark (clade: Acanthodii), but Ozarcus nests in a different node (Fig. 1) and shows how jaws developed from anterior gill arches. Despite the Early Carboniferous appearance of Ozarcus in the fossil record, the genesis of this taxon goes back to the mid-Silurian based on phylogenetic bracketing.
Ozarcus mapesae (Pradel et al. 2019; Early Carboniferous, 325 mya; AMNH FF20544) nests with Falcatus in the LRT, between Loganiella and Doliodus. The authors considered the palatoquadrate (pq) a single bone (Fig. 3) rather than the jugal, pterygoid, ectopterygoid, quadrate and lacrimal shown here (Fig. 2). The intertemporal anchors the hyomandibular here (eh) as in other vertebrates. The teeth are extremely tiny, much smaller than each tooth basin in the maxilla. The premaxilla is a tiny area between the ventrally opening incurrent nares. A ghosted prefrontal is shown framing the excurrent naris. The seemingly crushed narrowness of the skull was likely broader in life, especially around the gill basket.
Sister taxa, like Falcatus
(Fig. 4; another Early Carboniferous late survivor of an earlier Silurian radiation), helps us imagine what the post-crania of Ozarcus might look like. Falcatus also had a soft rostrum made of nasal bone precursors.
Lacking such a rostrum is Doliodus
(Fig. 5) from the Early Devonian. Rather than a tall narrow skull, as in Ozarcus, Doliodus had a wider skull with odd, two-pronged teeth. Such teeth represent an early experiment as toothed jaws were a new thing.
A basal elasmobranch still living with us today,
Chimaera, also has a tall narrow skull, like that of Ozarcus. Chimaera gives us a model for restoring what the post-crania of Ozarcus might have looked like.
Taxa in the LRT
document a gradual accumulation of derived traits. This is well illustrated with these related taxa, all of which look like one another, more so than any other included taxon. The tree topology has settled down now. New taxa drop in without upsetting the nesting of taxa already in the tree.
Pradel A, Maisey JG, Tafforeau P, Mapes RH and Mallant J 2014. A Palaeozoic shark with osteichthyan-like branchial arches. Nature 13185. doi:10.1038/nature13195e