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Phylogeny and Taxonomy of the Traversodontidae

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Book cover Early Evolutionary History of the Synapsida

Part of the book series: Vertebrate Paleobiology and Paleoanthropology ((VERT))

Abstract

We review the taxonomic history of traversodontid cynodont genera and species and the previous classifications and phylogenetic analyses of the group. 17 genera and 22 species are accepted as valid taxa within Traversodontidae. The phylogenetic relationships of traversodontids and other gomphodonts (including 6 trirachodontid species and Diademodon tetragonus) are analyzed based on 77 characters. Scalenodon angustifrons and (Andescynodon+Pascualgnathus) are found to be the most basal traversodontids. The three species of Scalenodon included in the analysis (Scalenodon angustifrons, S. attridgei, and S. hirschsoni) do not form a clade, supporting recognition of a separate genus for the latter two. A monophyletic Gomphodontosuchinae is recovered, as suggested by previous analyses. The distribution of traversodontids is discussed, with Gondwana and especially Africa suggested as the ancestral area for Gomphodontia and Traversodontidae.

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Acknowledgments

We thank Christian F. Kammerer, Agustín G. Martinelli, and an anonymous reviewer whose comments greatly improved this paper. The cooperation and hospitality of the staff of various museums and institutions greatly facilitated our comparative studies. We would like to thank Michael W. Maisch (GPIT), Sandra Chapman (NMHUK), Tom Kemp (Oxford), Ray Symonds (CAMZM), Johann Neveling (CGP), Jennifer Botha and Elize Butler (NMQR), Roger Smith and Sheena Kaal (SAM), Alejandro Kramarz and Agustín G. Martinelli (MACN), Ana Maria Ribeiro (MCN), Marcelo Reguero and Rosendo Pascual (MLP), Maria C. Malabarba (MCP), Guillermo F. Vega (PULR), Jaime E. Powell (PVL), Ricardo Martinez (PVSJ), Marina B. Soares and Cesar L. Schultz (UFRGS), John Flynn (AMNH), Olivier Rieppel, Elaine Zeiger, and William F. Simpson (FMNH), Charles R. Schaff and Wu Shaoyuan (MCZ), Hans-Dieter Sues and Matthew Carrano (USNM), James A. Hopson (University of Chicago), and Vince Schneider (NCSM). Financial support for this project was provided by the Chinese Academy of Sciences (KZCX2-YW BR-07), Columbia University through a Faculty Fellowship, the Climate Center of Lamont-Doherty Earth Observatory, and the Theodore Roosevelt Memorial Fund of AMNH. The Field Museum provided grants making research visits to Chicago possible. FA is supported by the National Research Foundation, South Africa.

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Authors and Affiliations

  1. Key Laboratory of Vertebrate Evolution and Human Origin of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, 100044, Beijing, People’s Republic of China

    Jun Liu

  2. Evolutionary Studies Institute, Private Bag 3, Wits, Johannesburg, 2050, South Africa

    Fernando Abdala

Authors
  1. Jun Liu
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  2. Fernando Abdala
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Corresponding author

Correspondence to Jun Liu .

Editor information

Editors and Affiliations

  1. Division of Paleontology and Richard Gilder, Graduate School American Museum of Natural History, New York, New York, USA

    Christian F. Kammerer

  2. Department of Geology, Field Museum, Chicago, Illinois, USA

    Kenneth D. Angielczyk

  3. Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt-Universität zu Berlin, Berlin, Germany

    Jörg Fröbisch

Appendices

Appendix 15.1: List of Morphological Characters

The following abbreviations are used to identify authors that previously used a particular character in data matrices: R, (Rowe 1988); M, (Martínez et al. 1996); F, (Flynn et al. 2000); H, (Hopson and Kitching 2001); A, (Abdala and Ribeiro 2003); Ab, (Abdala et al. 2006); SH, (Sidor and Hancox 2006); K, (Kammerer et al. 2008); S, (Sues and Hopson 2010); L, (Liu and Olsen 2010).

  1. 1.

    Adult maximum skull size: large (greater than 25 cm) (0), medium to small (1).

  2. 2.

    Snout (preorbit) in adults in relation to temporal region: longer (0), subequal (1), shorter (2) [Ab28].

  3. 3.

    Two side of temporal fenestra: divergent posteriorly (0), nearly parallel (1), bulge in the middle (2) [H39, Ab33].

  4. 4.

    Premaxilla forms posterior border incisive foramen: absent (0), present (1) [H1, Ab29].

  5. 5.

    Vomer exposure in incisive foramen (at anterior ends of maxillae on palate): present (0), absent (1) [M21].

  6. 6.

    Vomer: with (0) or without (1) vertical septum extending posteriorly beyond level of secondary palate [SH65].

  7. 7.

    Internarial bar: present (0), absent (1) [F5, A22, Ab21].

  8. 8.

    Parietal foramen in adults: present (0), absent (1) [H7, F6, A24, Ab23].

  9. 9.

    Ectopterygoid: present (0), absent (1) [H9, Ab30].

  10. 10.

    The posterior extension of secondary palate relative to anterior border of orbit: shorter (0), subequal (1), longer (2) [H15, S2].

  11. 11.

    Posterior extension of the jugal dorsally above the squamosal in the zygomatic arch: absent or with a small extension (0), well-developed (1) [F15, A26, Ab25].

  12. 12.

    The position of anterior root of the zygomatic arch relative to the ventral margin of the maxilla: nearly at same level or slightly higher (0), remarkable higher (1).

  13. 13.

    Zygomatic process of the jugal: little projected (0), conspicuously projected (1), absent (2), a ball-like process (3) [F16, H21, A25, Ab24].

  14. 14.

    Diameter of suborbital bar below center of orbit (anterior to suborbital process, where present): greater than 1/2 diameter of bar below posterior part of orbit (posterior to suborbital process) (0), less than 1/2 diameter of bar below posterior part of orbit (1) [S35].

  15. 15.

    Maxilla in the margin of the subtemporal fenestra: excluded (0), included (1) [Ab31].

  16. 16.

    Epipterygoid-quadrate contact: present (0), absent (1) [Ab32].

  17. 17.

    Frontal-epipterygoid contact: present (0), absent (1) [R39, H35].

  18. 18.

    Palatine: does not meet frontal (0), meets frontal but neither element contributes significantly to medial orbit wall (1) [H23].

  19. 19.

    Notch separating lambdoidal crest from zygomatic arch: shallow (0); deep, V-shaped (1) [H43].

  20. 20.

    Lower jaw symphysis as a chin-like process in adult: absent or little developed (0), well developed (1).

  21. 21.

    Dentary with sigmoid ventral curvature: absent (0), present (1) [S34].

  22. 22.

    Dentary angular process: not or very weakly projected posteriorly (0), projected posteriorly as distinct process (1) [A28, Ab27, S32].

  23. 23.

    Elongated mental foramen below postcanine tooth row and above coronoid ridge: absent (0), present (1) [S29].

  24. 24.

    Coronoid ridge anterior to masseteric fossa: absent to low (0), very strong, outturned (1) [S30].

  25. 25.

    Position of the upper canine in relation to paracanine fossa: posterolateral (0), lateral (1), anterolateral (2) [A6, Ab5, S1].

  26. 26.

    Diastema between upper incisors and canine: present (0), absent (1) [A3, Ab2].

  27. 27.

    Diastema between canine and maxillary postcanines in adult: short (0), long (1) [F1].

  28. 28.

    Diastema between canine and dentary postcanines: long (0), absent or very short (less than one tooth length) (1).

  29. 29.

    Maxillary labial platform lateral to the postcanine series: absent (0), present (1) [H77, A23, Ab22].

  30. 30.

    Posteromedial inclination of the last few upper gomphodont postcanines: absent or small (0), oblique (1) [F9, H72, A9, Ab8].

  31. 31.

    Axis of posterior part of maxillary tooth row: directed lateral to subtemporal fossa (0), directed towards center of fossa (1), directed toward medial rim of fossa (2) [H78, Ab36].

  32. 32.

    Maxillary tooth row extent relative to anterior margin of the subtemporal fossa in adult: anterior (0), at the same level (1), posterior (2).

  33. 33.

    Coronoid process of the mandible: covers the last postcanine (0), does not cover (1) [A27, Ab26].

  34. 34.

    Postcanine occlusion: absent (0), present (1) [Ab41].

  35. 35.

    Shearing planes between the outer surface of the main cusp of the lower and the inner surfaces of the main cusps of the uppers postcanines: present (0), absent (1) [Ab42].

  36. 36.

    Upper incisor number: four (0), three (1) [H53, F4, A1, Ab0].

  37. 37.

    Lower incisor number: three (0), two (1) [H54].

  38. 38.

    Incisor procumbency: (0) absent; (1) present [K43].

  39. 39.

    Incisor cutting margins: serrated (0), smoothly ridged (1), denticulated (2) [H55, Ab34].

  40. 40.

    Incisor size: small (0), enlarged (1) [F3, H56, A2, Ab1].

  41. 41.

    Upper canine size: large (0), reduced in size (1) [H57, A4, Ab3].

  42. 42.

    Lower canine size: large (0), reduced in size (1) [H58, A5, Ab4].

  43. 43.

    Canine serrations: present (0), absent (1) [H59, Ab35].

  44. 44.

    Postcanine tooth row in adults: formed by sectorial (0), conical, gomphodont and sectorial (1), gomphodont and sectorial (2), gomphodont (3) [H80, Ab37].

  45. 45.

    Overall morphology of the upper gomphodont postcanines in occlusive view: ovoid-ellipsoid (0), nearly rectangular (1), nearly triangular (2) [A7, Ab6].

  46. 46.

    Labial cingulum on anterior portion of the upper postcanines (external to the sectorial ridge): absent (0), present (1) [H61, A14, Ab13].

  47. 47.

    Posterior cingulum on upper postcanines: present (0), absent (1) [F7, A13, Ab12].

  48. 48.

    Shouldering in the posterior margin of upper postcanines: absent (0), slightly developed (1), well developed (2). [F2, A8, Ab7]

  49. 49.

    Anterior profile of principal labial cusp: convex (0), concave (1) [S31].

  50. 50.

    Number of cusps in the transverse crest of the upper postcanines: two (0), three or more (1) [F8, H63, A11, Ab10].

  51. 51.

    Central cusp of upper transverse row: midway between labial and lingual cusps (0), closer to lingual cusp (1) [H65, A12, Ab11].

  52. 52.

    Anterolabial accessory cusp on upper postcanines: one (0), absent (1), two or more (2) [H67].

  53. 53.

    Posterolabial accessory cusp on upper postcanines: present (0), absent (1) [H68].

  54. 54.

    Position of upper transverse cusp row on crown: central (0), anterior half of crown (1), at posterior part (2) [H64, A10, Ab9].

  55. 55.

    Distinct anterolingual accessory cusp on upper postcanines: absent (0), present (1) [H69, A15, Ab14].

  56. 56.

    Anterior cingulum in the upper postcanines: present (0), absent (1) [Ab39].

  57. 57.

    Anterior transverse crest on upper postcanines: absent or low (0), high (1) [F14, H70].

  58. 58.

    Lingual ridge on upper postcanines: absent (0), present (1) [F10, H71].

  59. 59.

    Overall morphology of the lower gomphodont postcanines in occlusal view: circular (0), ovoid-ellipsoid (1), quadrangular (2) [H62, A17, Ab16].

  60. 60.

    Transverse crest in lower postcanines: central (0), anterior (1) [A18, Ab17].

  61. 61.

    Number of cusps in the transverse crest of the lower postcanines: two (0), three or more (1) [H73, A19, Ab18].

  62. 62.

    Anterior cingulum in the lower postcanines: cuspules disposed on the entire margin (0), one or more cuspules located anterolabially (1), absent (2) [H74, A21, Ab20].

  63. 63.

    Posterior cingulum on the lower gomphodont postcanines: present (0), absent (1) [Ab38].

  64. 64.

    Size of the anterior cusps in the lower postcanines: labial lower than lingual (0), labial higher than lingual (1) [A20, Ab19].

  65. 65.

    Widest lower cusp in transverse row of lower postcanines: lingual (0), labial (1), middle (2) [F11, H76].

  66. 66.

    Anterolingual cusp of lower postcanines: nearly vertical (0); strongly inclined posteriorly (1).

  67. 67.

    Deep occlusal basins in the postcanines: absent (0), present (1) [H75, Ab40].

  68. 68.

    Anapophysis: absent (0), present (1) [L124].

  69. 69.

    Expanded costal plates on ribs: present (0), absent (1) [H82].

  70. 70.

    Lumbar costal plates with ridge overlapping preceding rib: present (0), absent (1) [H83].

  71. 71.

    Procoracoid in glenoid: present (0), barely present or absent (1) [H88].

  72. 72.

    The dorsal margin of the coracoid in medial view in relation to that of the procoracoid: shorter (0), equal or longer than (1).

  73. 73.

    Cranial margin of the procoracoid: convex (0), nearly straight (1), obviously concave (2).

  74. 74.

    Angle between ventral margin on anterior and posterior process of iliac blade: small (e.g., less than 140°) (0), large (1).

  75. 75.

    Length of anterior process of ilium anterior to acetabulum (relative to diameter of acetabulum): less than 1.5 (0), greater than 1.5 (1) [H94].

  76. 76.

    Dorsal profile of ilium: strongly convex (0), flat to concave (1) [H96].

  77. 77.

    The trochanter major position relative to the femoral head: distal (0), close, major part in same height (1).

Appendix 15.2: Character Matrix

Thrinaxodon

1020000000

0020010000

000000000?

0000?00010

0010???00?

-??-??????

???????101

0000000

Cynognathus

0000000000

0000001000

000000010?

0000?00000

0000???00?

-??-??????

???????000

0000000

Diademodon

0000000000

0010001010

0000000000

00(01)1000000

0001000000

-000?00000

000???0100

0001000

Beishanodon

0210?00101

002000??1?

????000?10

10???0?0??

0????0000?

??00?000??

??????0???

???????

Cricodon

1??0??0???

0???0????0

000000?010

1011100001

0002000001

1000?00010

100???0100

??????0

Langbergia

1110100001

001000?010

0000000000

1011100010

0002000001

0000?00010

100???0???

???????

Sinognathus

121??1?101

00200???10

0000000010

1011101010

00?2000001

0110??0010

10????0???

???????

Trirachodon

1110100(01)01

0010001010

0000000010

10(01)1100000

0002000001

(01)11(02)?00010

100???0100

000000?

JSM100

1????00??1

?0?0?????0

0000??0000

?0?1???0?0

0012100001

1110?00020

100???0???

???????

Andescynodon

11001101?1

00000???10

0000000010

2101000010

0003101000

-101?11021

011101110?

?111111

Arctotraversodon

0?????????

?????????1

0?11???0??

???10?0021

???3?01011

0?120?0?21

1212001???

???????

B . jeffersoni

112??101?2

002001??10

0111000010

2111000010

001220(01)111

1(02)12010121

(01)111(01)0101?

112???1

Dadadon

11101?01?1

?0300???1?

????110?11

21?100?010

0?13111111

10120101??

??????1???

???????

E . argentinus

0001111111

111111?110

1100211011

2201010111

011310121?

-00-111121

021111101?

1?11111

E . riograndensis

0001111111

111111??10

1100211?11

22?1010111

011310121?

-00-111121

021111101?

????11?

Gomphodontosuchus

????1?0??0

?1??0????0

1?00110011

2001000?10

111310111?

-??-011121

02111?1???

???????

L . drysdalli

101???00??

00110??010

1100000111

2101000000

0003010011

1(02)12000121

0101001101

0001110

L . sudamericana

1?????00?1

?001?????1

1100000111

2101000000

0003010011

1002000121

01010?1???

???????

M . ochagaviae

1211110112

002001?01?

?000110011

2(12)010000?0

0013101111

12120?0121

02?1101??1

???????

M . pascuali

1111110111

0020010010

000011001(01)

2(12)01000020

1113101111

1212000121

020010100?

1111111

Menadon

10111?0??0

11210???10

1100111011

2001000111

110310111?

-01-011121

021???101?

1?0111?

Pascualgnathus

110011?1?1

00000??010

0000000010

2101010010

0013000000

-101?11021

0?????1101

00?0011

Protuberum

101?111011

111111??1?

????211?11

22?101?111

1?131?1110

-0121111??

??????110?

???????

Santacruzodon

1?????0??0

?03??????0

1100000011

2101000020

??13201111

1212000121

0201011???

???????

S . angustifrons

110???00??

?0200???10

??00000010

2001000000

0003011011

111(02)010021

0100001???

???00?0

S . attridgei

???0110??1

????0?????

????000?11

21?101?010

0?13100011

10121011??

??????1???

???????

S . hirschoni

1??0110?11

?0?001?1?0

0?00000011

2101011011

1013100011

1012100121

0100001???

???????

Scalenodontoides

012???11?1

11?11???11

?10021101?

2201010111

?1?310121?

-00-?11121

0211111???

???????

Traversodon

1???1?00?1

?1110????1

1100000011

2101000010

0013110011

1?02000121

02?0?01?01

11????1

Nanogomphodon

??????????

??????????

??????????

???10?????

??????????

????????21

1101101???

???????

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Liu, J., Abdala, F. (2014). Phylogeny and Taxonomy of the Traversodontidae. In: Kammerer, C., Angielczyk, K., Fröbisch, J. (eds) Early Evolutionary History of the Synapsida. Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-6841-3_15

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