ALBERT

Description: The transition from water to land by the earliest tetrapods in the Devonian Period is seen as one of the greatest steps in evolution. However, little is understood concerning changes in brain morphology over this transition. Here, we determine the brain–braincase relationship in fishes and basal lissamphibians as a proxy to elucidate the changes that occurred over the fish–tetrapod transition. We investigate six basal extant sarcopterygians spanning coelacanths to salamanders ( Latimeria chalumnae, Neoceratodus, Protopterus aethiopicus, P. dolloi, Cynops, Ambystoma mexicanum ) using micro-CT and MRI and quantify the brain–braincase relationship in these extant taxa. Our results show that regions of lowest brain–endocast disparity are associated with regions of bony reinforcement directly adjacent to masticatory musculature for the mandible except in Neoceratodus and Latimeria . In Latimeria this deviation from the trend can be accounted for by the possession of an intracranial joint and basicranial muscles, whereas in Neoceratodus difference is attributed to dermal bones contributing to the overall neurocranial reinforcement. Besides Neoceratodus and Latimeria, regions of low brain–endocast disparity occur where there is less reinforcement away from high mandibular muscle mass, where the trigeminal nerve complex exits the braincase and where endolymphatic sacs occupy space between the brain and braincase wall. Despite basal tetrapods possessing reduced adductor muscle mass and a different biting mechanism to piscine sarcopterygians, regions of the neurocranium lacking osteological reinforcement in the basal tetrapods Lethiscus and Brachydectes broadly correspond to regions of high brain–endocast disparity seen in extant taxa.

Description: The fish-tetrapod transition (which incorporates the related fin-limb and water-land transitions) is celebrated as one of the most important junctions in vertebrate evolution. Sarcopterygian fishes (the “lobe-fins”) are today represented by lungfishes and coelacanths, but during the Paleozoic they were much more diverse. It was some of these sarcopterygians, a lineage of the tetrapodomorph fishes, that gave rise to tetrapods (terrestrial vertebrates with limbs bearing digits). This spectacular leap took place during the Devonian Period. Due to the nature of preservation, it is the hard parts of an animal’s body that are most likely to fossilize, while soft tissues such as muscular and brain tissues, typically fail to do so. Thus, our understanding of the adaptations of the hard skeletal structures of vertebrates is considerably greater than that of the soft tissue systems. Fortunately, the braincases of early vertebrates are often ossified and thereby have the potential to provide detailed morphological information. However, the correspondence between brain and endocast (an internal mold of the cavity) has historically been considered poor in most “lower” vertebrates and consequently neglected in such studies of brain evolution. Despite this, recent work documenting the spatial relationship in extant basal sarcopterygians (coelacanth, lungfish, axolotl, and salamander) has highlighted that this is not uniformly the case. Herein, we quantify and illustrate the brain-endocast relationship in four additional extant basal tetrapod exemplars: neobatrachian anurans (frogs) Breviceps poweri and Ceratophrys ornata; and gymnophionans (caecilians) Gegeneophis ramaswamii and Rhinatrema bivittatum. We show that anurans and caecilians appear to have brains that fill their endocasts to a similar degree to that of lungfishes and salamanders, but not coelacanth. Ceratophrys has considerably lower correspondence between the brain and endocast in the olfactory tract and mesencephalic regions, while Breviceps has low correspondence along its ventral endocranial margin. The brains of caecilians reflect their endocasts most closely (vol. ∼70%). The telencephalon is tightly fitted within the endocast in all four taxa. Our findings highlight the need to adequately assess the brain-endocast relationship in a broad range of vertebrates, in order to inform neural reconstructions of fossil taxa using the Extant Phylogenetic Bracket approach and future studies of brain evolution.