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Reappraisal of the Genus Tapirus (Perissodactyla, Tapiridae): Systematics and Phylogenetic Affinities of the South American Tapirs

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Abstract

The record of the genus Tapirus in South America is associated with the faunistic events of the Great American Biotic Interchange (GABI). The taxon is considered an immigrant of Holarctic origin. Although remains are scarce and incomplete during the Pleistocene, an analysis of these materials allowed us to consider valid seven fossil species : Tapirus tarijensis, T. cristatellus, T. greslebini, T. rioplatensis, T. oliverasi, T. mesopotamicus, and T. rondoniensis. A phylogenetic analysis was carried out in order to elucidate the relationships of the American fossil and extant species. Our result is consistent with a paraphyletic hypothesis for South American tapirs and suggests that a second dispersal event would have occurred from South America to North America, of a form closely related to T. cristatellus, resulting in the derived forms of North America.

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Acknowledgments

The authors thank Jorge I. Noriega, Mario A. Cozzuol, and Marco Aurélio G. de França for their valuable comments on drafts of the manuscript and support. We thank Alejandro G. Kramarz and David Flores (MACN), Cástor Cartelle (PUCMinas), Daniel A. García López (PVL), João A. de Oliveira (MNRJ), Márcia Martins (MCN), Mariano Merino (MLP), Mario de Vivo (MZUSP), Miguel Sant’Anna (UNIR), and Richard C. Hulbert (UF) for access to material under their care. We thank M. Griselda Gottardi (CICYTTP-Conicet) who prepared the fossil materials. We also thank Alessandra D. Boos for language revision, and two anonymous referees who helped improve the manuscript. This study was partially funded by Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina, and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Brazil. The first author thanks the Florida Museum of Natural History for the travel grant awarded. Contribution to the projects: PIP 886; PICT-ANPCYT 392 and CNPq/PROSUL490299/2008-3.

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Authors and Affiliations

  1. Programa de Pós-Graduação em Geociências, Universidade Federal do Rio Grande do Sul, Porto Alegre, RS, Brazil

    Elizete Celestino Holanda

  2. Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción de Diamante-CONICET, Diamante, Entre Ríos, Argentina

    Brenda Soledad Ferrero

Authors
  1. Elizete Celestino Holanda
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  2. Brenda Soledad Ferrero
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Correspondence to Elizete Celestino Holanda.

Appendices

Appendix 1. Description of Characters Used for Phylogenetic Analysis

The characters and character states are adapted from the character set originally developed by Hulbert and Wallace (2005), with phylogenetically uninformative characters deleted. Original numbering scheme of characters is retained in parentheses.

  1. 0.

    width of sagittal crest in adult specimens: (0) narrow; (1) broad [modified from Hulbert 1995: #2].

  2. 1 (3).

    temporal crests meet to form sagittal crest: (0) at or posterior to the frontoparietal suture; (1) anterior to the frontoparietal suture.

  3. 2 (4).

    dorsal table of frontal: (0) relatively narrow or small; (1) relatively broad.

  4. 3 (5).

    frontal inflation: (0) weak or absent; (1) strong.

  5. 4 (6).

    nasal-frontal lateral profile: (0) nasals and frontals approximately on the same plane; (1) nasals notably stepped down from frontals.

  6. 5 (7).

    size of interparietal bone compared to the parietal: (0) large, interparietal occupying more than 1/3 length of parietal in sagittal plane; (1) small, interparietal occupying less than 1/3 length of parietal in sagittal plane [modified];

  7. 6 (8).

    shape of interparietal bone: (0) typically polygonal (hexagonal or diamond shaped); (1) typically triangular.

  8. 7 (9).

    interparietal fusion with occipital: (0) occurs early in ontogeny (before loss of DP4); (1) occurs late in ontogeny (after eruption of P4).

  9. 8 (10).

    nasal length: (0) long (longer than 1.5 times the width of combined nasals); (1) short (shorter than 1.5 times the width of combined nasals).

  10. 9 (11).

    anterolateral margin of nasal: (0) relatively straight; (1) distinctly concave.

  11. 10 (14).

    fossa for meatal diverticulum on nasal: (0) shallow and without distinct margins; (1) deep and with distinct margins.

  12. 11 (15).

    fossa for meatal diverticulum on posterior dorsal surface of nasal: (0) not extensive, does not near midline; (1) very extensive, approaches within a few mm of midline.

  13. 12 (16).

    development of fossa for meatal diverticulum on dorsal table of frontal: (0) very limited; (1) broad exposure with distinct posterior margin.

  14. 13 (17).

    supraorbital groove for nasal diverticulum: (0) broad and shallow; (1) narrow and deep.

  15. 14 (18).

    posterodorsal process of maxilla widely exposed dorsal to orbit, forming base of trough for meatal diverticulum: (0) yes; (1) no.

  16. 15 (20).

    orientation of lambdoidal crests in adults: (0) mostly posteriorly, little or no outward flair; (1) posterolateral orientation, with notable lateral flair.

  17. 16 (22).

    location of infraorbital foramen relative to cheek teeth: (0) dorsal to P4; (1) dorsal to P3 or P2.

  18. 17 (24).

    anteromedial process of maxilla: (0) absent; (1) present.

  19. 18 (25).

    lateral exposure of anteromedial process of maxilla: (0) maxilla well exposed in lateral view dorsal to premaxilla; (1) maxilla covered by premaxilla, not visible in lateral view or barely so.

  20. 19 (26).

    dorsal maxillary flange: (0) absent; (1) present [modified].

  21. 20 (28).

    width of maxillary bar between infraorbital foramen and lacrimal: (0) narrow, less than 5 mm; (1) wide, more than 5 mm.

  22. 21 (29).

    shape of lacrimal: (0) narrow, much taller than long; (1) broad, about as long as it is high.

  23. 22 (30).

    facial surface of lacrimal: (0) flat or convex; (1) concave.

  24. 23 (31).

    anterior lacrimal process(es): (0) absent or very weak; (1) present, well developed.

  25. 24 (32).

    posterior (preorbital) process of lacrimal: (0) absent; (1) present.

  26. 25 (33).

    posterior process of lacrimal: (0) broad and flat; (1) slender, pointed or knobby.

  27. 26 (34).

    typical number of lacrimal foramen: (0) two; (1) one.

  28. 27 (35).

    lacrimal foramen visible in lateral view: (0) yes, at least one; (1) no, not visible in lateral view.

  29. 28 (36).

    incisive foramen: (0) terminates posteriorly about midway between canine and P1; (1) terminates posteriorly at P1 or more posterior.

  30. 29 (37).

    relative diastema length: (0) short (ldl/p2m3L < 0.40); (1) long (ldl/p2m3L ≥0.40) [ldl = lower diastema length; p2m3L = length from the anterior end of p2 to the posterior end of m3]. [modified]

  31. 30 (38).

    relative location of mental foramen: (0) anterior to the p2; (1) directly ventral to the p2.

  32. 31 (39).

    orientation of anterior margin of ascending ramus of mandible in lateral view: (0) projects vertically and posteriorly, not anteriorly (does not overlie the m3); (1) projects anteriorly as well as vertically below the coronoid process, often lies dorsal to m3.

  33. 32 (40).

    depth of horizontal ramus below m3: (0) shallow (m3 depth <18 % mandible length); (1) deep (m3 depth > 18 % of mandible length) [modified from Hulbert 2010: #40].

  34. 33 (41).

    relative height of crown in cheek teeth: (0) short, relatively brachydont; (1) taller.

  35. 34 (42).

    morphology and size of I3: (0) I3 incisiform, about same size as I2; (1) I3 caniniform, much larger than I2 [Colbert 2005: #3].

  36. 35 (45).

    P2 AW/PW ratio (anterior width/posterior width): (0) on average, less than or equal to 0.85; (1) on average, more than 0.85.

  37. 36 (46).

    P2 lingual cingulum: (0) complete; (1) absent or “broken” [modified from Colbert 2005: #29]

  38. 37 (47).

    transverse lophs on P2-P4 and p2-p4: (0) poorly separated; (1) well separated.

  39. 38 (48).

    P2 protoloph: (0) does not reach ectoloph; (1) just reaches base of ectoloph.

  40. 39 (49).

    P3 protoloph: (0) does not reach ectoloph or just reaches base of ectoloph; (1) merges midway or higher onto ectoloph [modified].

  41. 40 (51).

    P2 metaloph: (0) does not reach ectoloph or just reaches base of ectoloph; (1) merges midway or higher onto ectoloph [modified].

  42. 41 (54).

    P3 and P4 protocone and hypocone: (0) closely appressed and merged with wear; (1) widely separated.

  43. 42 (55).

    parastyle development in P3-M3: (0) moderate, parastyle height less than 50 % of paracone height; (1) strong, parastyle height more than 50 % of paracone height, but parastyle smaller than paracone; (2) very strong, parastyle height equal to paracone height [modified].

  44. 43 (56).

    labial cingulum on posterior half of upper cheek teeth: (0) present on half or more of P3-M3; (1) absent or very rare in P3-M3.

  45. 44 (67).

    i1 morphology and size: (0) i1 larger than i2, not procumbent or slightly procumbent; (1) i1 much larger than i2, very procumbent and spatulate. [modified].

  46. 45 (69).

    length of p2 relative to p3: (0) short, (p2 L)/(p3 L) less than 1.1; (1) long, (p2L)/(p3L) ratio more than 1.1.

  47. 46 (70).

    p2 hypoconid and entoconid: (0) not joined by complete hypolophid; (1) connected by hypolophid.

  48. 47 (72).

    relative height of unworn protolophid and hypolophid (or hypoconid/entoconid if no hypolophid) in premolars: (0) protolophid distinctly taller than hypolophid; (1) the two are approximately equal in height.

  49. 48 (73).

    cristid obliqua on p3: (0) strong, blocks interlophid valley labially; (1) very weak or absent.

  50. 49 (74).

    cristid obliqua on p4: (0) strong, blocks interlophid valley labially; (1) very weak or absent.

  51. 50 (78).

    relative length of limbs (comparing length of femur and dentary): (0) relatively long limbs (femur greatest length longer than that of dentary); (1) relatively short limbs (femur length less than or equal to that of dentary).

Appendix 2. Taxon-Character Matrix

Table 4 Character numbers in top row correspond to those in Appendix 1. “?” indicates an unknown or non applicable character state. a = 0 or 1. Numbers in bold correspond to character states revised and coded different from the original matrix of Hulbert and Wallace (2005) or Ferrero and Noriega (2007)
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Holanda, E.C., Ferrero, B.S. Reappraisal of the Genus Tapirus (Perissodactyla, Tapiridae): Systematics and Phylogenetic Affinities of the South American Tapirs. J Mammal Evol 20, 33–44 (2013). https://doi.org/10.1007/s10914-012-9196-z

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