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A complex brain is central to the success of backboned animals. However, direct evidence bearing on vertebrate brain evolution comes almost exclusively from extant species, leaving substantial knowledge gaps. Although rare, soft-tissue preservation in fossils can yield unique insights on patterns of neuroanatomical evolution. Paleontological evidence from an exceptionally preserved Pennsylvanian (ca. 318 Ma) actinopterygian, Coccocephalus, calls into question prior interpretations of ancestral actinopterygian brain conditions. However, ordering and timing of major evolutionary innovations such as an everted telencephalon, modified meningeal tissues, and hypothalamic inferior lobes remain unclear. Here we report two distinct actinopterygian morphotypes from the latest Carboniferous-earliest Permian (~299 Ma) of Brazil that show extensive soft-tissue preservation of brains, cranial nerves, eyes and potential cardiovascular tissues. These fossils corroborate inferences drawn from Coccocephalus, while adding new information about neuroanatomical evolution. Skeletal features indicate that one of these Brazilian morphotypes is more closely related to living actinopterygians than the other, which is also reflected in soft-tissue features. Significantly, the more crownward morphotype shows a key neuroanatomical feature of extant actinopterygians–an everted telencephalon–that is absent in the other morphotype and Coccocephalus. All preserved Paleozoic actinopterygian brains show broad similarities including an invaginated cerebellum, hypothalamus inferior lobes, and a small forebrain. In each case, preserved brains are substantially smaller than the enclosing cranial chamber. The neuroanatomical similarities shared by this grade of Permo-Carboniferous actinopterygians reflect probable primitive conditions for actinopterygians, providing a revised model for interpreting brain evolution in a major branch of the vertebrate tree of life. 

Abstract Though Paleozoic ray‐finned fishes are considered to be morphologically conservative, we report a novel mode of fang accommodation (i.e., the fitting of fangs inside the jaw) in the Permian actinopterygian †Brazilichthys macrognathus, whereby the teeth of the lower jaw insert into fenestrae of the upper jaw. To better understand how fishes have accommodated lower jaw fangs through geologic time, we synthesize the multitude of ways living and extinct osteichthyans have housed large mandibular dentition. While the precise structure of fang accommodation seen in †Brazilichthyshas not been reported in any other osteichthyans, alternate strategies of upper jaw fenestration to fit mandibular fangs are present in some extant ray‐finned fishes—the needlejawsAcestrorhynchusand the gars of the genusLepisosteus. Notably, out of our survey, only the two aforementioned neopterygians bear upper jaw fenestration for the accommodation of mandibular fangs. We implicate the kinetic jaws of neopterygians in this trend, whereby large mandibular fangs are more easily fit between the multitude of upper jaw and palatal bones. The restricted space available in early osteichthyan jaws may have led to a proliferation of novel ways to accommodate large dentition. We recommend a greater survey of Paleozoic actinopterygian jaw morphology, in light of these results and other recent reevaluations of jaw structure in early fossil ray‐fins.