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Exoskeleton and Systematics: A Historical Problem in the Classification of Glyptodonts

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Abstract

The glyptodonts (Mammalia: Cingulata) are characterized by an exoskeleton comprising most notably an armored tail and an immobile dorsal carapace formed by a large number of osteoderms. In 1889, Florentino Ameghino published the first phylogenetic scenario for the glyptodonts, based on the sequential application of two transformation series related to the morphology of the tail armor and carapace osteoderms. From the early to mid 1900s, several authors used Ameghino’s transformation series subordinated to a model of evolution in which derived glyptodont groups had arisen independently from separate pre-middle Miocene ancestors. This approach, in which the morphological states of Ameghino’s series were considered analogous rather than homologous, provided different phylogenetic scenarios and the paraphyletic classification still in use. Two recent cladistic analyses based on cranial and postcranial (including caudal tube) characters support the monophyly of glyptodonts and suggest novel intra-clade relationships. However, neither analysis included the classic osteoderm characters used by earlier authors. Therefore, we propose new osteoderm and carapace characters and evaluate their performance in a new cladistic analysis. We found that: a) some osteoderm characters used by earlier authors to support ancestor-descendent hypotheses are in fact fully homoplastic autapomorphies (e.g., multiplication of the number of rows of peripheral figures); b) characters previously believed to have originated independently in several groups (e.g., presence of caudal tube) are synapomorphies at a wider hierarchical level; c) some ancestor–descendant pre-cladistic hypotheses are incompatible with the topology and synapomorphy distribution obtained; and d) there is no reason to favor exoskeletal characters in glyptodont systematics.

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Acknowledgements

We are grateful to Timothy Gaudin and François Pujos for inviting us to participate in this volume, and to T. Gaudin and two anonymous reviewers for their interesting suggestions that improved the manuscript. This is a contribution to the grants, PICT 0143, UNLP N 647 and PIP-CONICET 1054 to Sergio F. Vizcaíno and UNLu CDD-CD 281-09 to Juan Carlos Fernicola. Kleberson O. Porpino would like to acknowledge CNPq for financial support (project 401825/2010-8).

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Authors and Affiliations

  1. CONICET, División Paleontología de Vertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,”, Av. Ángel Gallardo 470 1405DJR, Ciudad Autónoma de Buenos Aires, Argentina

    J. C. Fernicola

  2. Departamento de Ciencias Básicas, Universidad Nacional de Luján, Ruta 5 y Av. Constitución, 6700, Luján, Buenos Aires, Argentina

    J. C. Fernicola

  3. Departamento de Ciências Biológicas, Universidade do Estado do Rio Grande do Norte, Rua Antônio Campos s/n, Mossoró, 59610-090, Rio Grande do Norte, Brazil

    K. O. Porpino

Authors
  1. J. C. Fernicola
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  2. K. O. Porpino
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Correspondence to J. C. Fernicola.

Appendices

Appendix I

Table 1 Specimens examined and references consulted for comparative study and cladistic analysis
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Appendix II

List of new characters and character states added to the matrix of Porpino et al. (2010) and used in the cladistic analysis. Characters marked with an asterisk (*) are ordered.

  • 01. External surface ornamentation of osteoderms: with figures defined by sulci (0); with a polygonal central area delimited by a dorsal thickening (1); without figures, smooth and perforated by large canals (2); without figures, presenting numerous small foramina associated with poorly developed sulci (3).

  • 02*. Main figure defined by a principal sulcus and peripheral figures: near the posterior edge and peripheral figures absent or poorly developed at the posterior border (0); roughly central, surrounded by a single row of well-developed peripheral figures (1); roughly central, surrounded by at least two single rows of well-developed peripheral figures (2).

  • 03. Outline of main figure defined by sulcus: circular or subcircular (0); roughly oblong (1).

  • 04. Transverse mobile band(s): complete (extending from side to side of the carapace) (0); incomplete (limited to the ventrolateral border of the anterior region) or absent (1).

Appendix III

Table 2 The scoring of the new osteoderm and carapace characters for the 18 taxa used in the cladistic analysis. States marked with N are inapplicable
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Fernicola, J.C., Porpino, K.O. Exoskeleton and Systematics: A Historical Problem in the Classification of Glyptodonts. J Mammal Evol 19, 171–183 (2012). https://doi.org/10.1007/s10914-012-9186-1

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