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A revision of the fossil genus Phanomys Ameghino, 1887 (Rodentia, Hystricognathi, Cavioidea) from the early Miocene of Patagonia (Argentina) and the acquisition of euhypsodonty in Cavioidea sensu stricto

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Abstract

Cavioidea sensu stricto (Cavioidea s.s.) is one of the most divergent lineages within the South American Hystricognathi, and is currently represented by cavies, maras (Caviidae), and capybaras (Hydrochoeridae). Caviids and hydrochoerids have been interpreted as forming the crown group, whereas the stem group of Cavioidea s.s. is formed by “Eocardiidae”, a paraphyletic group recorded in the late Oligocene to middle Miocene, mainly in Patagonia. One of the most interesting features of Cavioidea s.s. is the record of evolution of hypsodonty, which develops relatively slowly compared to other groups of caviomorphs, enabling this process to be followed in the fossil record. Phanomys is recorded in Patagonia during the late early Miocene, and since its description this genus has been considered close to euhypsodont genera. The objective of this paper is to: (1) report new material of Phanomys; (2) re-describe the two species of the genus; (3) determine the phylogenetic position of Phanomys among Cavioidea s.s. by morphological cladistic analysis; and (4) explore the sequence of appearance of characters related to the origin of euhypsodonty. The new materials assigned to Phanomys mixtus are the first mandibular fragments and palate known, yielding valuable morphological, ontogenetic, and phylogenetic information. Phanomys vetulus is recognized as a valid species, although better material is needed for a more robust definition. The phylogenetic analysis shows that Phanomys is the sister group of euhypsodont Cavioidea s.s. The evolutionary history of Cavioidea s.s. demonstrates that character states previously thought to correlate with the degree of hypsodonty (e.g. absence of fossettes/ids) do not all appear at the same time during the evolution of the group. Although the evolutionary trend in Cavioidea s.s. shows progressively increasing hypsodonty, the morphological changes inferred from this phylogenetic analysis indicate that hypsodonty and other character states were temporally decoupled during the evolution of the group during the Oligocene and early Miocene.

Kurzfassung

Die Cavioidea sensu stricto repräsentieren eine der divergierenden Linien innerhalb der su?damerikanischen Hystricognathi und sind zurzeit durch Meerschweinchen, Maras (Caviidae), und Wasserschweine vertreten. Caviiden und Hydrochoeriden wurden als die Kronen-Gruppe der Cavioidea s.s. interpretiert, während die “Eocardiidae”, eine paraphyletische Gruppe aus dem späten Oligozän bis mittleren Miozän aus Patagonien, als Stamm-Gruppe dargestellt wurden. Eine der interessantesten Merkmalsentwicklungen von Cavioidea s.s. ist die Evolution der Hypsodontie, die sich im Vergleich zu anderen Gruppen von Caviomorphen relativ langsam entwickelte; dies ermöglicht es, diesen Prozess bei Fossilien zu erkennen. Phanomys ist aus dem Ende des frühesten Miozän von Patagonien bekannt, und seit ihrer Erstbeschreibung wurde diese Gattung in die Nähe von euhypsodonten Taxa gestellt. Das Ziel dieses Beitrags ist es, neues Material von Phanomys vorzustellen, die beiden Arten dieser Gattung neu zu beschreiben, ihre phylogenetische Stellung innerhalb der Caviodea s.s. durch eine kladistische Analyse mit morphologischen Merkmalen zu untersuchen und die Sequenz des Merkmalserwerbes von mit Euhypsodontie verbundener Merkmale zu untersuchen. Bei dem neuen Material handelt es sich um den ersten Rest des Unterkiefers und des Gaumens von Phanomys mixtus, und diese Reste ergeben wichtige morphologische, ontogenetische und auch phylogenetische Informationen. Bei Phanomys vetulus handelt sich um eine gültige Art, obwohl besseres Material für eine befriedigende Definition nötig ist. Die phylogenetische Analyse zeigt, dass Phanomys die Schwestergruppe der euhypsodonten Cavioidea s.s. darstellt. Die Evolution der Cavioidea s.s. zeigt, dass die Merkmale, die bisher in direktem Zusammenhang mit dem Grad der Hypsodontie gestellt wurden (z.B. Abwesenheit von fossettes/ids) nicht zur gleichen Zeit während der Entwicklung der Gruppe erscheinen. Obwohl die evolutionäre Entwicklung bei den Caviodea s.s. einen Trend zu zunehmender Hypsodontie aufweist, zeigen die morphologischen Veränderungen, basierend auf dieser phylogenetischen Hypothese, dass diese Merkmale und der Erwerb der Hypsodenties in der Evolution der Gruppe im Oligozän und frühen Miozän zeitlich entkoppelt waren.

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Acknowledgments

We thank C. Deschamps (MLP), for improving the English version, and L. Reiner (MEF), for laboratory work. We thank Michelle Arnal (MACN) for improving the German zusammenfassung version. Access to SEM lab was possible thanks to ALUAR Aluminio Argentino SAIC and the help of Mr J. Groizard, and we thank M. Arnal, C. Vieytes and C. Deschamps for taking photos of MACN SC 3450. We thank the curators A. Kramarz (MACN), E. Ruigómez (MPEF-PV), M. Reguero (MLP-PV), D. Verzi (MLP), D. Romero (MMP), J. Flynn and J. Meng (AMNH), Christopher Norris and D. Brinkman (YPM PU), and K. Wellspring (ACM) for access to material in their care. We thank S. Vizcaíno and S. Bargo for access to material in their care. We are especially thankful to D. Pol for his aid in the phylogenetic analysis. Oliver Rauhut, Darin Croft, and A. Kramarz provided critical comments during the review process that greatly enhanced the quality of this manuscript. Fieldwork at Río Jeinemeni was supported by grants from the National Science Foundation to R. Kay. Comparisons with cavioid taxa for the phylogenetic analysis were made possible thanks to the Collection Study Grant (AMNH), an Ostrom Fund Grant (YPM, USA), PICT 38112 (M.G. Vucetich), and Fundación Egidio Feruglio.

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  1. Museo Paleontológico Egidio Feruglio, Avenida Fontana 140, U9100GYO, Trelew, Chubut, Argentina

    María E. Pérez

  2. Facultad de Ciencias Naturales y Museo, División Paleontología Vertebrados, Universidad Nacional de La Plata, Paseo del Bosque S/No, B1900FWA, Buenos Aires, Argentina

    María G. Vucetich

  3. CONICET, Buenos Aires, Argentina

    María E. Pérez & María G. Vucetich

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  1. María E. Pérez
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Correspondence to María E. Pérez.

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Appendix 1

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Table 3   
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Appendix 2

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Table 4 Character-taxon matrix (Pérez and Vucetich 2011) used for phylogenetic analysis was re-scored by Phanomys mixtus and Phanomys vetulus
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Pérez, M.E., Vucetich, M.G. A revision of the fossil genus Phanomys Ameghino, 1887 (Rodentia, Hystricognathi, Cavioidea) from the early Miocene of Patagonia (Argentina) and the acquisition of euhypsodonty in Cavioidea sensu stricto. Paläontol Z 86, 187–204 (2012). https://doi.org/10.1007/s12542-011-0120-2

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