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New information about Irajatherium hernandezi Martinelli, Bonaparte, Schultz & Rubert 2005 (Eucynodontia, Tritheledontidae) from the upper triassic (Caturrita Formation, Paraná Basin) of Brazil

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Abstract

In this contribution, new specimens of the tritheledontid eucynodont Irajatherium hernandezi, from the Late Triassic (Caturrita Formation) of southern Brazil, are analyzed. The new material provides significant information about incisor count, canine size and shape, basicranial morphology, and other previously unknown aspects of skull and dentition. A cladistic analysis with inclusion of the new data supports the assignment of Irajatherium to Tritheledontidae, basal to Chalimininae and Pachygenelinae. Previously unknown characters of Irajatherium revealed by the new material include: the presence of three lower incisors; the first lower incisor is enlarged; the presence of large upper and lower canines with deep paracanine fossa on the maxilla; almost complete upper and lower postcanine tooth row with a pattern similar to that of other tritheledontids (e.g. Pachygenelus and Chaliminia); there is a conspicuous crest on the inner surface of the maxilla for the attachment of the inferred maxillary turbinates; partially confluent jugular foramen and fenestra rotunda within the jugular fossa, separated by a finger-like projection of the posterolateral wall of the opisthotic; and hypoglossal foramina located outside the jugular fossa. Irajatherium is a key taxon for understanding the early evolution of ictidosaurs, a group of cynodonts closely related to mammaliaforms, during the cynodont–mammal transition from the Late Triassic to Early Jurassic.

Kurzfassung

In dieser Arbeit werden neue Exemplare des tritheledontiden Eucynodonten Irajatherium hernandezi Martinelli et al. 2005 aus der Späten Trias (Caturrita Formation) vorgestellt. Das neue Material enthüllt wichtige Informationen zur Anzahl der Incisiven, der Grösse und Form der Canini, der Morphologie des Basicraniums und anderen Aspekte von Schädel und Bezahnung. Eine cladistische Analyse mit Berücksichtigung der neuen Daten bestätigt die Zuordnung von Irajatherium zu den Tritheledontidae, welche basal der Chalimininae und Pachygenelinae stehen. Unter den vorher unbekannten Merkmale von Irajatherium sind drei untere Incisiven mit einem vergrößerten ersten Incisivus, untere und obere Canini mit einer tiefen Paracaninus-Fossa am Maxillare; eine fast komplette obere und untere postcanine Zahnreihe mit einem Bezahnungsmuster vergleichbar mit dem anderer Tritheledontiden (z.B. Pachygenelus und Chaliminia); eine auffällige, als Ansatzfläche für die Turbinalia dienende Crista an der Innenseite of the Maxilla; partiell vereinigte Jugular-Foramina und eine Fenestra rotunda in der Jugular-fossa, welche durch einen fingerartigen Vorsprung der posterolateralen Wandung des Opistothicums voneinander getrennt werden; sowie außerhalb der Jugular-Fossa liegene hypoglossale Foramina. Irajatherium ist ein Schlüsseltaxon zum Verständnis der frühen Evolution der Ictidosauriden, eine mit den Mammaliformen nahe verwandte Cynodontier-Gruppe aus der oberen Trias und dem unteren Jura.

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Abbreviations

UFRGS PV-T:

Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil, Paleovertebrate Collection.

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Acknowledgments

We thank Luiz Flávio Lopes for the photographs of several specimens and CNPq for financial support (TVO). The suggestions of the reviewers Fernando Abdala and Zhe-Xi Luo and the editor Oliver Rauhut improved the quality of this paper.

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Appendices

Appendix 1: Score for Irajatherium hernandezi in the data matrix of Martinelli & Rougier (2007)

  • Irajatherium

  • ?11?1?1002110?????100011101210???0110?????0????1???????????????????1000??1?1?????0????0111???

Changes in the score for Irajatherium hernandezi presented in this paper compared with the data matrix of Martinelli & Rougier (2007):

  • Character 2. Lower incisor number: from ? to 1 (three incisors).

  • Character 3. Some incisor enlarged: from ? to 1 (present).

  • Character 24. Sectorial postcanines with the major axis anteromedial-posterolaterally oriented, passing the distal portion of an anterior tooth medial to the mesial portion of the next tooth: from 0 to 1 (present).

  • Character 35. Mandibular symphysis: from ? to 1 (unfused).

  • Character 36. Postdentary bones: from ? to 1 (reduced to narrow rod lying in postdentary trough).

  • Character 68. Trigeminal nerve (V2 and V3) exit: from ? to 1 (foramen located entirely on the prootic, or on the anterior lamina of the petrosal).

  • Character 69. Prootic and ophistotic: from ? to 0 (unfused).

  • Character 70. Petrosal promontorium: from ? to 0 (absent).

  • Character 71. Fenestra ovalis: from ? to 0 (with thickened ring).

  • Character 74. Perilymphatic foramen separation from jugular foramen: from ? to 1 (partially separated by finger-like projection from postero-lateral wall of jugular foramen).

  • Character 76. Hypoglossal foramen: from ? to 1 (separated from the jugular fossa).

Appendix 2: Diagnoses of clades (numbered as in Fig. 9; some clades are not diagnosed here). An asterisk (*) indicates ambiguous character-states

  • Clade 4. Ictidosauria (sensu Martinelli & Rougier, 2007): presence of some enlarged incisors (Ch. 3), upper incisor 2 large and the others small (Ch. 4), lower incisor 1 large and the other small (Ch. 5), premaxilla forming the posterior border of incisive foramen (Ch. 39)*, length of palatine longer relative to maxilla in secondary palate (Ch. 47).

  • Clade 5: Tritheledontidae (sensu Martinelli & Rougier, 2007): dominant central bulbous main cusp on upper postcanines (Ch. 11), upper posterior postcanines with cusp B and C buccally displaced and bulbous, prominent cusp A (Ch. 12), lower middle and posterior postcanines with four cusps aligned decreasing in size posteriorly (Ch. 19), posterior portion of secondary palate almost at the level of the tip of upper postcanine teeth, forming a deep groove between hard palate and tooth row (Ch. 48).

  • Unnamed clade. Pachygenelinae (Clade 6) plus Chalimininae (Clade 7): single roots of postcanines (Ch. 25), tooth row parallel to sub-parallel from the axial plane of the cranium (Ch. 30), maxillary buccal shelf overhanging tooth row (Ch. 43).

  • Clade 6. Pachygenelinae (sensu Martinelli & Rougier, 2007): upper postcanines with buccal cingulum (Ch. 9)*, narrow upper postcanines with lingual cingulum (Ch. 10)*.

  • Clade 7. Chalimininae (sensu Martinelli & Rougier, 2007): more than 11 upper postcanine teeth in adult (Ch. 8)*, absence of imbrincated sectorial postcanines (Ch. 24)*.

  • Tritheledon plus Diarthrognathus: absence of upper posterior postcanines with cusp B and C buccally displaced and bulbous, prominent cusp A (Ch. 12)*.

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de Oliveira, T.V., Martinelli, A.G. & Soares, M.B. New information about Irajatherium hernandezi Martinelli, Bonaparte, Schultz & Rubert 2005 (Eucynodontia, Tritheledontidae) from the upper triassic (Caturrita Formation, Paraná Basin) of Brazil. Paläontol Z 85, 67–82 (2011). https://doi.org/10.1007/s12542-010-0078-5

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