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Study of a digital cranial endocast of the non-mammaliaform cynodont Brasilitherium riograndensis (Later Triassic, Brazil) and its relevance to the evolution of the mammalian brain

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Abstract

A digital cranial endocast of the specimen UFRGS-PV-1043-T, Brasilitherium riograndensis, was obtained from high-resolution computed tomography (μCT) scan images. This taxon is a small cynodont from the Late Triassic of Brazil, and has been used as the sister-group of the mammaliaforms in cladistic analyses. The digital endocast of UFRGS-PV-1043-T is mostly complete, allowing the description and collection of accurate linear and volumetric measurements, which were taken and compared with other non-mammaliaform cynodonts. Impressions of vessels were observed in the inner walls of the braincase. Despite the lack of a cribiform plate and the presence of a wide orbital vacuity, the endocast of Brasilitherium shows olfactory bulb casts that are relatively larger than in other non-mammaliaform cynodonts, suggesting a pattern of gradual increase in size and improvement of the olfactory sense for these structures toward the mammalian condition. The cerebral hemispheres are elongated and clearly divided by a median sulcus. The parafloccular casts are well defined, and their position corresponds to the maximum width of the endocast. In the ventral view, a large hypophyseal cast and a wide opening for the cavum epiptericum are evident. The encephalization quotient (EQ) calculated for Brasilitherium is greater than the range of EQs reported for most non-mammaliaform cynodonts (although it may be similar to that of some taxa, according to the equation used to estimate their body masses), but it is smaller than that of the mammaliaforms and mammals. A slighter increase in the brain size of Brasilitherium compared with other non-mammaliaform cynodonts was observed, along with a more significant increase in the size of the olfactory bulbs. This study supports the proposition of an early evolution of the mammalian brain associated with selective pressures for better sensorial acuity, especially regarding improved olfaction, which began with small Triassic mammaliamorphs.

Kurzfassung

Mittels μCT-Daten wurde ein virtueller Gehirnausguss von Brasilitherium riograndensis (UFRGS-PV-1043-T) erstellt. Bei diesem Taxon handelt es sich um einen kleinen Cynodontier aus der späten Trias von Brasilien, der in cladistischen Analysen die Schwestergruppe der Mammaliaformes repräsentiert. Der fast vollständige virtuelle Gehirnausguss von UFRGS-PV-1043-T ermöglicht die Beschreibung sowie genaueste lineare und volumetrische Messungen im Vergleich mit anderen nicht-mammaliaformen Cynodontier. Am inneren Schädeldach befinden sich einige Abdrücke von Blutgefässen. Obwohl eine Lamina cribrosa fehlt und die Augenhöhlen sehr groß sind, weist der virtuelle Ausguss von Brasilitherium relativ größere Bulbi olfactorii auf als bei anderen nicht-mammaliaformen Cynodontiern. Somit lässt sich eine graduelle Größenzunahme sowie Verbesserung des Riechsinns hin zu einem säugerähnlichen Zustand beobachten. Die Großhirnhälften sind verlängert und deutlich durch einen medianen Sulcus voneinander getrennt. Die Paraflocculusausgüsse sind deutlich ausgeprägt und ihre Lage korrespondiert mit der maximalen Breite des gesamten Gehirnausgusses. In ventraler Ansicht ist ein prominenter Ausguss der Hypophyse sowie eine große Öffnung in das Cavum epiptericum zu erkennen. Der für Brasilitherium berechnete Enzephalisationsquotient (EQ) ist größer als die Spannbreite der EQs der meisten anderen nicht-mammaliaformen Cynodontier. Aufgrund der Schätzung des Körpergewichts von Brasilitherium könnte dessen EQ jedoch dem bestimmter Taxa durchaus entsprechen; er ist jedoch sicherlich geringer als bei Mammaliaformes und Mammalia. Brasilitherium zeigt eine geringere Größenzunahme des Gehirns im Vergleich zu anderen nicht-mammaliaformen Cynodontiern, mit wesentlich deutlicherer Größenzunahme der Bulbi olfactorii. Diese Studie unterstützt die Annahme einer frühen Evolution des Säugergehirns assoziiert mit der Selektion erhöhter sensorischer Schärfe, insbesondere der Verbesserung der Riechleistung, die bereits bei triassischen kleinen Mammaliamorpha auftritt.

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Acknowledgments

The fieldwork for data collection was supported by the National Geographic Society and the Ligabue Foundation of Venice (Italy). The specimen was prepared by José Fernando Bonaparte (UFRGS/NGS) and Agustin Martinelli (Sección Paleontología de Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”), and was taken to Germany by JFB to be scanned at the Hochschule Aalen through a project submitted to the Alexander von Humboldt Foundation with the participation of Wolfgang Maier (Institut für Evolution und Ökologie, Eberhard-Karls-Universität Tübingen).

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  1. Laboratório de Paleovertebrados, Departamento de Paleontologia e Estratigrafia, Instituto de Geociências, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves, Porto Alegre, RS, 9500, Brazil

    Pablo Gusmão Rodrigues & Cesar Leandro Schultz

  2. Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Nussallee 8, 53115, Bonn, Germany

    Irina Ruf

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Rodrigues, P.G., Ruf, I. & Schultz, C.L. Study of a digital cranial endocast of the non-mammaliaform cynodont Brasilitherium riograndensis (Later Triassic, Brazil) and its relevance to the evolution of the mammalian brain. Paläontol Z 88, 329–352 (2014). https://doi.org/10.1007/s12542-013-0200-6

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