Abstract

Most authors have identified two rapid increases in relative brain size (encephalization quotient, EQ) in cetacean evolution: first at the origin of the modern suborders (odontocetes and mysticetes) around the Eocene-Oligocene transition, and a second at the origin of the delphinoid odontocetes during the middle Miocene. We explore how methods used to estimate brain and body mass alter this perceived timing and rate of cetacean EQ evolution. We provide new data on modern mammals (mysticetes, odontocetes, and terrestrial artiodactyls) and show that brain mass and endocranial volume scale allometrically, and that endocranial volume is not a direct proxy for brain mass. We demonstrate that inconsistencies in the methods used to estimate body size across the Eocene-Oligocene boundary have caused a spurious pattern in earlier relative brain size studies. Instead, we employ a single method, using occipital condyle width as a skeletal proxy for body mass using a new dataset of extant cetaceans, to clarify this pattern. We suggest that cetacean relative brain size is most accurately portrayed using EQs based on the scaling coefficients as observed in the closely related terrestrial artiodactyls. Finally, we include additional data for an Eocene whale, raising the sample size of Eocene archaeocetes to seven. Our analysis of fossil cetacean EQ is different from previous works which had shown that a sudden increase in EQ coincided with the origin of odontocetes at the Eocene-Oligocene boundary. Instead, our data show that brain size increased at the origin of basilosaurids, 5 million years before the Eocene-Oligocene transition, and we do not observe a significant increase in relative brain size at the origin of odontocetes.

Highlights

  • Cetaceans such as the sperm and killer whales have brains exceeding nine kilograms [1, 2], larger than any other species on the planet, living or extinct

  • We re-evaluate the existing data on brain and body size in fossil cetaceans, with a focus on the Eocene, and we present the cetacean data alongside their terrestrial artiodactyl relatives

  • These data, in addition to previously published values (S1 Table in S2 File) are used to examine the relationship between brain mass and endocranial volume in extant taxa, the quantification of which forms the basis of our brain mass estimates in fossil cetaceans and terrestrial artiodactyls

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Summary

Introduction

Cetaceans such as the sperm and killer whales have brains exceeding nine kilograms [1, 2], larger than any other species on the planet, living or extinct. Scaled for body size, the largest cetacean brains are eclipsed only by those of the genus Homo [3, 4]. Most studies comparing brain sizes use an index called the encephalization quotient (EQ) to accommodate the fact that animals with larger bodies tend to have proportionally larger brains [2, 4,5,6,7,8,9,10,11,12]. The EQ value indicates how much larger (EQ>1) or smaller (EQ

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