Supplementary Material The early Miocene balaenid Morenocetus parvus



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Supplementary Material

The early Miocene balaenid Morenocetus parvus from Patagonia (Argentina) and the evolution of right whales

Mónica R. Buono, Marta S. Fernández, Mario A. Cozzuol, Jose I. Cuitiño, and Erich M. G. Fitzgerald.


Contents

Institutional abbreviations

List of the specimens studied for the anatomical and phylogenetic analyses

List of characters

Figure S1

Figure S2

Supplementary Table S1

Supplementary Table S2

Supplementary Table S3 Supplementary references


Institutional abbreviations. AMNH, American Museum of natural History, New York, USA; ChM, The Charleston Museum, Charleston, USA; CNPMAMM, Laboratorio de Mamíferos Marinos, Centro Nacional Patagónico, Puerto Madryn, Argentina; FMNH,  Field Museum of Natural History , Chicago, USA; HUES, Hokkaido University of Education Sapporo, Sapporo, Hokkaido, Japan; IRSNM, Institute Royal des Sciences Naturelles, Bruxelles, Belgium; LACM, Natural History Museum of Los Angeles County, Los Angeles, California, USA; MACN, Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina; MFM, Mizunami Fossil Museum, Gifu, Japan; MLP, Museo de Ciencias Naturales de La Plata, La Plata, Argentina; MSNT; Museo di Storia Naturale e del Territorio, Universita di Pisa, Pisa, Italy; MNHN, Museum National d‘Histoire Naturelle, Paris, France; NMB, Natuurmuseum Brabant, Tilburg, The Netherlands; NMNS, National Museum of Nature and Science, Japan; NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand; NMV, Museums Victoria, Melbourne, Australia; OU, Geology Museum, Department of Geology, University of Otago, Dunedin, New Zealand; SFM, Shinshushinmachi Fossil Museum, Shinshushinmachi, Nagano Prefecture, Japan; UCMP, Museum of Paleontology, University of California, Berkeley, California, USA; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; SFM, Shinshushinmachi Fossil Museum, Shinshushinmachi, Nagano Prefecture, Japan; SMNS, Staatliches Museum für Naturkunde, Stuttgart, Germany; SBAER, Soprintendenza per i Beni Archeologici dell’Emilia Romagna, Italy; ZMUC: Zoological Museum of the Natural History Museum, Copenhagen, Denmark.
List of the specimens studied for the anatomical and phylogenetic analyses:

The data matrix includes 43 taxa: one outgroup (the archaeocete Zyghoriza kochii), two odontocete species (Physeter macrocephalus and Waipatia maerewhenua), and 40 mysticete species (31 extinct and 9 extant taxa) representing a broad sample of mysticete diversity. We decided not to include Balaenula balaenopsis in our analysis because the taxonomic status of this taxon needs to be carefully revised because there are serious doubts that all of the skeletal elements referred by Van Beneden (1872) actually represent one individual, let alone one taxon.

The list includes the specimens used in the phylogenetic and comparative analyses. The taxa marked with an asterisk were studied in person by MRB. Other specimens were analysed from the literature (the references are specified) and/or from photographs provided by colleagues.

Taxon

Collection number

References

*Zygorhiza kochii

USNM 11962




*Waipatia maerewhenua

cast of the holotype OU22095




*Physeter macrocephalus

USNM 35315

USNM 253051






Janjucetus hunderi

NMV P216929

Fitzgerald, 2006

Mammalodon colliveri

NMV P199986

Fitzgerald, 2010

*Aetiocetus cotylalveus

USNM 25210




Aetiocetus weltoni

UCMP 12290

Berta & Demeré, 2008

Fucaia goedertorum

LACM 5930




Eomysticetus whitmorei

ChM PV4253

Sanders & Barnes, 2002

Micromysticetus rothauseni

ChM PV4844

Sanders & Barnes, 2002

*Morenocetus parvus

MLP 5-11

MLP 5-15


Cabrera 1926

*Balaena mysticetus

*USNM 257513

*USNM 259000

*USNM 63300

LACM 54479 (photographs provided by F. Marx)

LACM 54485, (photographs provided by F. Marx)

ZMUC CN1 (photographs provided by F. Marx)






Balaena montalionis

MSNT I 12357 (photographs provided by F. Marx)

Capellini 1904; Bisconti 2000, 2003

*Balaena ricei

USNM 22553

Westgate & Whitmore Jr. 2002

Balaenella brachyrhynus

NMB 42001 (photographs provided by F. Marx)

Bisconti 2005

Balaenula sp.

HUES 10003 (photographs provided by F. Marx)

Kimura 2009

Balaenula astensis

MSNT I12555 (photographs provided by F. Marx)

Bisconti 2002, 2003; Pilleri, 1987


Eubalaena ianitrix

IRSNM CtM 879a-f

Plisnier-Ladame & Quinet, 1969; Bisconti et al., 2017

Eubalaena shinshuensis

SFM CV-0024 (photographs provided by M. Churchill)

Kimura 2009

*Eubalaena australis

CNPMAMM 748

CNPMAMM 746 CNPMAMM 774 CNPMAMM 742

MLP 1508

MACN 54.119

USNM 484901

USNM 267612

USNM 484899

USNM 2676129

NMNZ 2239

OU 226


Schaeff et al., 1999; Schaeff & Hamilton, 1999

*Eubalaena glacialis

*USNM 23077

*USNM 301637

*USNM 500860

*USNM 504343

*USNM 504886

ZMUC CN2 (photographs provided by F. Marx)






*Eubalaena japonica

*USNM 16434

*USNM 16435

NMNS M33436 (photographs provided by F. Marx)





Peripolocetus vexillifer

CAS 4370 (photographs provided by F. Marx)

Kellog, 1931

*Caperea marginata

USNM 340

NMNZ MM 2235







Miocaperea pulchra

SMNS 46978

Bisconti, 2012

Herpetocetus morrowi

UCMP 124950

Adli et al., 2014

*Herpetocetus transatlanticus

*USNM 182962

USNM 299653

USNM 312543


Whitmore & Barnes, 2008

Isanacetus laticephalus

MFM 18004

MFM 28501



Kimura & Ozawa, 2002

Cetotherium rathkii

PIN 1840/1 (photographs provided by F. Marx)

Brandt, 1843

*Diorocetus hiatus

USNM 16783




*Nannocetus eremus

USNM 244236




*Pelocetus calvertensis

USNM 11976




*Aglaocetus moreni

*MLP 5-14

FMNH P13407






*Aglaocetus patulus

USNM 23690

USNM 13472






Mauicetus parki

OU 11573

Benham, 1937

Piscobalaena nana

MNHN 1616

MNHN 1617

MNHN 259





Eschrichtioides gastaldii

MRSN PU13802

Bisconti, 2008

*Eschrichtius robustus

USNM 364973

AMNH 181374






Archaeobalaenoptera castriarquati

SBAER 240536

Bisconti, 2007

*Balaenoptera bonaerensis

CNPMAMM 750

MACN 20522






*Balaenoptera musculus

USNM 124326

MLP1199





*Megaptera miocaena

USNM 23690




*Megaptera novaengliae

USNM 13656

USNM 21492






List of characters.

257 morphological characters were used in the phylogenetic analyses. The source of characters was principally from Marx & Fordyce (2015), plus other sources (Bisconti 2005, Churchill et al., 2012; characters 15, 255, 256 and 257). Characters 27-37, 67, 96, 226, 246 and 252 were excluded because they represent apomorphies of taxa excluded from our analysis. Characters 2, 21, and 42 were modified from Marx & Fordyce (2015) (see below). In addition the character codings of some taxa were changed from the original matrix (which is specified in each case, see below). Some characters were treated as ordered following Marx & Fordyce (2015).



Cranium

1. Length of rostral portion of maxilla anterior to antorbital notch: less than bizygomatic width (0); equal to or greater than bizygomatic width (1); more than one and a half times the bizygomatic width (2). ORDERED



Note: The original coding of Balaenella was changed from “0” to “1”

2. Portion of rostrum anterior to nasals in lateral view: below the level of the supraoccipital (0); raised to the level of the supraoccipital (1); raised above the level of the supraoccipital (2) (modified from Marx and Fordyce, 2015; Churchill et al 2012).



Note: the original coding of Balaena was changed from “1” to “2”.

3. Lateral edge of maxilla in cross section: forms an angle of more than 45 degrees (0); lateral edge is flattened with an angle of less than 45 degrees (1).

4. Lateral border of maxilla anterior to antorbital notch (or homologous point on rostrum) in dorsal view: concave (0); straight or slightly convex (1); broadly convex (2). ORDERED

5. Transverse width of maxilla at midpoint: distinctly less than that of the premaxilla (0); roughly equal to or up to twice the width of the premaxilla (1); more than twice the width of the premaxilla (2). ORDERED

6. Premaxilla in dorsal view: widens at anterior end (0); portion anterior to nasal opening narrows or remains the same width anteriorly (1).

7. Premaxilla adjacent to and anterior to narial fossa: elevated above the maxilla and forming a distinct lateral face (0); continuous or nearly continuous with the maxilla (1).

8. Premaxilla adjacent to and at posterior edge of nasal opening: does not clearly overhang maxilla (0); premaxilla overhangs maxilla (1).

9. Anterior portions of premaxillae: firmly contact each other (0); premaxillae are separated or only loosely contact along their entire length (1).

10. Suture between maxilla and premaxilla on rostrum: firmly articulated (0); loose (1).

11. Antorbital process: absent (0); present and defined by a steep face clearly separating the posterolateral corner of the maxilla from its more anterior rostral portion (1); present as a distinct anterior projection lateral to antorbital notch (2).

12. Anterior border of supraorbital process lateral to ascending process of the maxilla: bordered by lacrimal and maxilla (0); bordered by lacrimal only (1); as state 0, but with the antorbital process of the maxilla and the anterior border of the supraorbital process separated by a basin (2); as state 0 but with maxilla overriding the anteriormost border of the supraorbital process (3).

13. Distinct pocket between the ascending process of the maxilla dorsally and the supraorbital process ventrally: absent (0); present (1).

14. Lateral process of maxilla (sensu Marx, Lambert & Uhen, 2016): absent (0); present and clearly distinct from ascending process of maxilla (1); present and confluent with ascending process of maxilla (2).

15. Lateral process of the maxilla: directed posterolaterally at an oblique angle to the anteroposterior axis of the cranium; directed laterally, nearly perpendicular to the anteroposterior axis of the cranium (Churchill, Berta & Deméré, 2012; Bisconti, 2005).

16. Posterior portion of palatal surface of maxilla: flattened or slightly concave (0); medial portion of maxilla forms a longitudinal keel bordered laterally by a shallow longitudinal trough (1); medial portion forms a keel without any adjacent trough (2).

17. Palatal surface of anterior part of rostrum: flat or gently concave (0); bears pronounced longitudinal keel formed by the vomer and the medial edges of the maxillae (1).

18. Exposure of premaxilla on palate: exposed along at least one third of the medial border of the maxilla (0); limited in extent to less than one third of the medial border of the maxilla (1).

19. Palatal window exposing vomer: absent (0); present (1); narrow and variable exposure of vomer along most or all of the midline of the rostrum (2); vomer broadly exposed along the midline of the rostrum (3).

20. Palatal nutrient foramina and sulci: absent (0); present (1).

21. Outline of suture between maxillae, vomer and palatines: roughly straight transversely or bowed anteriorly (0); forms a posteriorly pointing V shape (1); anterior margins of palatines form two separate and posteriorly pointing U shapes (2) (modified from Marx & Fordyce, 2015).

22. Anteriormost point of palatine: located in line with or posterior to the level of the antorbital notch or equivalent point on rostrum (0); located anterior to the level of the antorbital notch (1).

23. Anterior portion of palatine distinctly concave transversely and forming a sharp median crest: absent (0); present (1).

24. Anterior edge of nasal fossa: located in posterior three quarters of rostrum (0); located in anterior quarter of rostrum (1).

25. Facial portion of rostrum in lateral view: step-like (0); straight (1); concave (2). ORDERED

26. Rostrum shape: width at antorbital notches or equivalent point on rostrum less than 80% the length of the rostrum, as measured from its tip to the antorbital notches (0); width at antorbital notches or equivalent point more than 80% the length of the rostrum (1).

27. Teeth in adult individuals: present (0); absent or vestigial (1).

28. Skull length about one third or more of total body length: absent (0); present (1).

29. Cranial asymmetry: present (0); absent (1).

30. Diameter of orbit as measured between the distalmost points of the preorbital and postorbital processes: less than 25% of bizygomatic width (0); 25% or more (1).

31. Anterior edge of supraorbital process lateral to ascending process of maxilla with the skull in dorsal view: oriented transversely or pointing anteriorly (0); pointing posteriorly (1); linguiform and tapering to a point (2).



Note: The coding of Balaenula astensis and Balaenula sp. was changed from “1” to “0”.

32. Outline of anterior edge of supraorbital process in dorsal view: roughly straight or concave (0); distinctly sinusoidal (1).

33. Transverse width of anterior edge of supraorbital process lateral to ascending process of maxilla: longer than or equal to the combined transverse width of the adjacent rostral bones, as measured from the sagittal plane to the lateral border of the ascending process of the maxilla (0); shorter than the combined transverse width of the adjacent rostral bones (1).

34. Posterior border of supraorbital process in dorsal view: concave (0); straight (1).

35. Supraorbital process of frontal in anterior view: horizontal or nearly horizontal (0); gradually slopes away lateroventrally from the skull vertex (1); as state 1 but with the lateral portion of the supraorbital being nearly horizontal, thus causing the latter to appear concave in anterior view (2); abruptly depressed to a level noticeably below the vertex, with the lateral skull wall above the supraorbital formed by both parietal and frontal (3). ORDERED

36. Anterior and posterior borders of supraorbital process in dorsal view: roughly parallel or converging medially (0); converging laterally (1).

37. Width of supraorbital process as measured in a straight line from the lateralmost point of the postorbital process to the intertemporal constriction: equal to or shorter than the anteroposterior length of the supraorbital process above the orbit (0); up to twice the length above the orbit (1); more than twice the length above the orbit (2). ORDERED

38. Postorbital process in dorsal view: oriented posteriorly (0); oriented laterally (1); oriented posterolaterally (2); short and not markedly projecting in any direction (3).

39. Postorbital process in lateral view: pointed or rounded (0); forms and anteroposteriorly elongate triangle with a flattened posterior face (1).

40. Orbital rim of supraorbital process of frontal in lateral view: dorsoventrally thin (0); thickened with a flat lateral surface (1); thickened with a rounded lateral surface (2).



Note: The coding of Morenocetus was changed from “0” to “1” and Balaenula astensis from “2” “0”.

41. Position of anteriormost point of supraorbital process in dorsal view: in line with the posterior extremity of the nasals or passing through the nasals (0); at the same level as the anterior extremity of the nasals (1); anterior to the anterior extremity of the nasals (2). ORDERED



Note: the coding of Morenocetus was changed from “0” to “?” because the estimated most anterior point of the supraorbital process of frontal is posterior to the posterior margin of the nasals. In addition, the coding of Eubalaena australis was changed from “-“ to “0”.

42. Position of the dorsal margin of the orbit (in lateral view, with skull resting on a horizontal surface): above or in line with the vertex of the skull (0); located roughly a half of distance between the vertex of the skull and the ventral surface of the postglenoid process (1) located a 2/3 the vertical distance between the vertex of the skull and the ventral surface of the postglenoid process (2); located a 1/3 the vertical distance between the vertex of the skull and the ventral surface of the postglenoid process (3) (modified from Marx & Fordyce, 2015).

43. Lacrimal in dorsal view: situated entirely lateral to the ascending process of the maxilla (0); lacrimal extends medially and separates the lateral corner of the ascending process from the more anterior portion of the maxilla (1).

44. Contact of jugal with zygomatic process of squamosal: the two bones overlap dorsoventrally (0); little or no overlap (1).

45. Anteriormost portion of jugal broadly underlapped by maxilla: absent (0); present (1).

46. Optic canal in ventral view: ventrally open (0); medial portion is enclosed by anterior and/or posterior bony laminae (1).

47. Postorbital ridge along medial portion of optic canal: absent or anteroposteriorly thin, with the optic canal running adjacent to the posterior border of the supraorbital process (0); well developed and thickened, thus resulting in the displacement of the optic canal away from the posterior border of the supraorbital process (1).

48. Maxillary infraorbital plate (sensu Marx, Lambert & Uhen, 2016): absent (0); present (1).



Note: We changed the coding of Morenocetus from “1” to “?”

49. Maxillary window (sensu Marx, Lambert & Uhen, 2016) originating from posterior border of infraorbital plate: absent (0); present (1).

50. Anteromedial corner of supraorbital process extending to a point medial to antorbital notch: absent (0); present (1).

Note: We changed the coding of Morenocetus from “0” to “?”.

51. Preorbital region of frontal in lateral view: thickened compared to more central portions of the orbit (0); dorsoventrally flat (1).

52. Enlarged dorsal infraorbital foramen on ascending process of maxilla, opening into a posterodorsally directed sulcus: absent (0); present (1).

53. Premaxillary sac fossa: absent (0); present (1).

54. Premaxillary foramen: absent (0); present (1).

55. Suture between maxilla and frontal: contact between the bones is straight or maxilla overrides anteromedial corner of the frontal (0); maxilla overrides half or more of the frontal (1).

56. Lateral borders of ascending process of maxilla: lateral edges convergent with the process tapering to a point (0); lateral edges parallel or divergent posteriorly (1).

57. Triangular wedge of frontal separating ascending process of maxilla from nasal orpremaxilla: absent (0); present (1).

58. Posterior ends of ascending processes of maxillae in dorsal view: separated by either frontals or both nasals and premaxillae (0); converging towards the midline and separated by nasals only (1); contact each other medially (2). ORDERED

59. Relative position of posteriormost edge of ascending process of maxilla in dorsal view: approximately in transverse line with, or posterior to, posterior edge of nasal (0); anterior to posterior edge of nasal (1).

60. Shape of posterior border of ascending process of maxilla: pointed or rounded (0); squared off (1).

61. Lateral profile of cranium along exposure of parietals on vertex: dorsal edge of parietal ascends steeply towards posterior edge of skull at an angle of 10 degrees or more, measured relative to the lateral edge of rostrum (0); dorsal edge of parietal is low to flat with an angle of less than 10 degrees (1).

62. Length of nasal relative to bizygomatic width: less than 50% of bizygomatic width (0); more than 50% of bizygomatic width (1).

63. Lateral margins of nasal: parallel (0); posteriorly convergent (1).

64. Anterior margins of nasals: roughly straight or U-shaped (0); form a distinct, posteriorly pointing W-shape (1); with point on midline and a gap on each side between premaxilla and nasal (2);

65. Dorsal surface of nasals: flattened (0); developed into a sagittal keel (1).



Note: The coding of Morenocetus was changed from “1” to “?”; the nasals are not preserved in this taxon.

66. Separation of posterior portions of nasals along sagittal plane by narial process of frontal: present (0); absent (1).



Note: the narial process is clearly developed in Eubalaena glacialis, but is variably present in specimens of Eubalaena australis (it is coded as polymorphic).

67. Zygomatic process of squamosal and exoccipital in dorsal view: clearly separated by an angle (0); posterior border of zygomatic process and lateral edge of exoccipital are confluent, forming a continuous or nearly continuous lateral skull border (1).

68. Orbitotemporal crest: positioned along posterior border of supraorbital process, with the origin of the temporal muscle facing posteriorly or posteroventrally (0); absent or positioned on the dorsal surface of the supraorbital process, with the origin of the temporal muscle facing posterodorsally or dorsally (1).

69. Area enclosed by orbitotemporal crest on supraorbital process of frontal: forms less than half of the dorsal surface of the supraorbital process (0); covers half or more of the dorsal surface of the supraorbital process (1).

70. Outline and orientation of orbitotemporal crest: subparallel to posterior border of supraorbital process (0); distal half oriented distinctly posterolaterally and approaching the posterolateral corner of the supraorbital process (1); as state 1, but with the crest terminating halfway along the posterior border of the supraorbital process (2); as state 2, but with the crest being distinctly U-shaped (3).

71. Shape of temporal fossa: longer anteroposteriorly than wide transversely, or as wide as long (0); wider than long (1).

72. Intertemporal constriction: longer anteroposteriorly than wide transversely (0); as state 1, but with the temporal fossa forming a large parasagittal oval (1); wider transversely than long anteroposteriorly (2).

73. Exposure of frontal on skull vertex: broadly exposed (0); anteroposteriorly compressed or absent (1).

74. Parietal and interparietal: anteriormost point located no further forward than postorbital process (0); anteriormost point in line with supraorbital process (1).

75. Outline of fronto-parietal suture: straight or lobate (0); frontals projects posteriorly along the sagittal plane and separate the left and right parietal anteriorly (1); highly irregular (2).



Note: we changed the coding from “?” to “0” in Morenocetus because the frontoparietal suture is evident in the dorsal view of the skull.

76. Parietal in lateral view: as long or longer anteroposteriorly than high dorsoventrally (0); higher dorsoventrally than long anteroposteriorly (1).

77. Spreading of anterolateral portion of parietal on to posteromedial corner of supraorbital process of frontal: absent (0); present (1).

78. Anteriormost point of parietal and interparietal: more posterior than the posterior border of the ascending process of the maxilla (0); more anterior than or in line with the posterior border of the ascending process of the maxilla (1).

79. Anteriormost point of supraoccipital in dorsal view: located posterior to or in line with the anterior border of the squamosal fossa (0); in line with temporal fossa, but posterior to the apex of the zygomatic process of the squamosal (1); in line with or located anterior to the level of the apex of the zygomatic process of the squamosal (2); in line with the anterior half or anterior edge of the supraorbital process (3). ORDERED

Note: we changed the coding of Morenocetus from “2” to “3” according with our proposed dorsal orientation of the skull.

80. Anteroposterior position of posterior apex of nuchal crest: posterior to the occipital condyle (0); anterior to or in line with the posteriormost point of the occipital condyle (1).

81. Mediolateral position of posterior apex of nuchal crest: aligned with the medial half or halfway point of the temporal fossa (0); approaching the level of the lateral border of the temp oral fossa (1).

82. Distinct tubercle at junction of parieto-squamosal suture and supraoccipital: absent (0); present (1).

83. Exposure of alisphenoid within or at ventral border of temporal fossa: exposed on temporal wall of skull and contributing to orbital fissure (0); alisphenoid separated from orbital fissure or not exposed on temporal skull wall (1).

84. Postparietal foramen located at junction of parietal and squamosal: absent (0); present (1).

85. Zygomatic process of squamosal dorsoventrally expanded in lateral view: absent (0); present, with the zygomatic process being distinctly higher dorsoventrally than wide transversely (1).

Note: the coding of Morenocetus was changed from “o, 1” to “1”. The zygomatic process is well preserved in the referred specimen of Morenocetus, and has a similar shape to that of extant balaenids.

86. Orientation of zygomatic process of squamosal in dorsal view: directed anteromedially (0); directed anteriorly or slightly anterolaterally (1); directed anterolaterally (2). ORDERED

87. Zygomatic process of squamosal in lateral or ventral view: tapering anteriorly (0);expanded anteriorly, thus forming a central constriction (1).

88. Twisting of zygomatic process of squamosal: absent (0); present, with the zygomatic process being partially twisted clockwise on the left and anticlockwise on the right (1); as state 1, but with the process twisted almost 90 degrees (2).

89. Position of apex of zygomatic process of squamosal: situated entirely posterior to the postorbital process (0); closely apposed to the postorbital process, or situated ventral to the latter (1).

90. Apex of zygomatic process of squamosal deflected anteroventrally: absent (0); present (1).



Note: the coding of Morenocetus was changed from “0” to “1” because, as is evident in the referred specimen MLP 5-15, the zygomatic process has a ventral orientation.

91. Supramastoid crest of zygomatic process of squamosal (skull in lateral view): present (0); absent (1).



Note: The coding of Balaenula spp. was changed from “?” to “1”

92. Size of squamosal including zygomatic and postglenoid processes: longer anteroposteriorly than high dorsoventrally, or about as high as long (0); distinctly higher than long (1).

93. Parieto-squamosal suture shaped like a crest or ridge: absent or low (0); present and distinctly elevated (1).

Note: The coding of Morenocetus was changed from “0” to “1” (a distinct crest is evident in MLP 5-15)

94. Squamosal prominence (sensu Fordyce & Marx, 2013): present as a projection on the crest delimiting the lateral or posterolateral edge of the squamosal fossa (0); absent (1).

95. Transverse width of squamosal lateral to exoccipital: width equal to or greater than 15% of the distance between the sagittal plane and the lateral edge of the exoccipital (0); exposed portion of squamosal is less than 15% of that distance (1).

96. Length of squamosal fossa relative to maximum transverse width of temporal fossa, as measured in a straight line from the posteriormost point of the temporal fossa to the posteriormost point of the nuchal crest: length of squamosal fossa is three quarters the width of the temporal fossa or longer (0); length of squamosal fossa is less than three quarters the width of the temporal fossa (1).

97. Squamosal cleft (sensu Marx, Lambert & Uhen, 2016): absent (0); present (1).

98. Squamosal crease: absent (0); present (1).

99. Paired tubercles on supraoccipital: absent (0); limited to low ridges forming the lateral edges of a medial fossa (1); present (2). ORDERED

Note: the coding of Morenocetus was changed from “0” to “1” because when the skull is viewed dorsally low ridges are visible delimiting a medial longitudinal fossa.

100. Lateral edge of supraoccipital in dorsal view: convex (0); straight (1); concave (2); sigmoidal (3).

101. Anterior border of supraoccipital shield: rounded or pointed (0); squared (1).

102. Overall outline of supraoccipital in dorsal view: rounded (0); triangular (1).

103. Anterior half of dorsal surface of supraoccipital: concave (0); flat or convex (1).

104. External occipital crest: absent or faint (0); restricted to anterior half of supraoccipital shield (1); present and running all the way along the supraoccipital shield (2). ORDERED

105. Tip of postglenoid process in lateral view: curving anteriorly (0); pointing ventrally (1); pointing posteriorly (2).

106. Ventral edge of postglenoid process in lateral view: approximately in line with or dorsal to the ventral edge of the exoccipital (0); extending well ventral to the ventral edge of the exoccipital (1).

107. Orientation of postglenoid process in posterior view: ventrolateral (0); ventral (1); ventromedial (2). ORDERED

108. Outline of postglenoid process in anterior or posterior view: parabolic (0); as state 0, but with lateral and medial edges parallel or concave (1); as state 1, but distinctly wider transversely than high dorsoventrally (2); triangular (3); trapezoidal, with a ventrally directed medial border (4).

109. Twisting of postglenoid process in ventral view: absent (0); twisted clockwise on the left side and anticlockwise on the right side so that the glenoid cavity faces anteromedially (1).

110. Position of base of postglenoid process in ventral or posterior view: in line with the lateral edge of the skull (0); shifted away medially from the lateral edge of the skull (1).

111. Medial border of postglenoid process in ventral view: confluent with more medial portion of squamosal (0); offset from remainder of squamosal by a distinct ridge (1).

112. Choanal margin of palatine in ventral view: absent (0); straight or convex (1); concave (2); forms a longitudinal notch (3).

113. Pterygoid in ventral view: partially or entirely exposed (0); palatine almost completely covers pterygoid and extends on to the hamular process (1).

114. Anteriormost point of pterygoid sinus fossa: located anterior to foramen pseudovale (0); approximately in line with anterior edge of foramen pseudovale (1); located posterior to anterior edge of foramen pseudovale (2). ORDERED

115. Dorsal lamina of pterygoid: absent or restricted to anteromedial quarter of pterygoid sinus fossa (0); present and covering half or more of ventral exposure of alisphenoid within pterygoid sinus fossa (1).

116. Shape of pterygoid hamulus: finger-like (0); expanded into a dorsoventrally flattened plate flooring the pterygoid sinus fossa (1); triangular and wing-like (2); reduced in size or almost absent (3).

117. Position of pterygoid hamulus in ventral view: located directly adjacent to the sagittal plane and almost contacting each other (0); well separated from each other (1).

118. Position of foramen pseudovale: foramen located within squamosal or between squamosal and pterygoid, and opening anterolaterally or laterally (0); as state 0, but with foramen opening posteriorly (1); foramen lies within pterygoid (2).

119. Foramen pseudovale raised above more lateral portions of squamosal in ventral view: absent (0); present (1).

120. Fossa on squamosal for reception of sigmoid process of tympanic bulla: present (0); absent or poorly defined (1).

121. Base of postglenoid process in ventral view: in transverse line with or located posterior to the posterior half of the tympanic bulla (0); in transverse line with or located anterior to the anterior half of the tympanic bulla (1); in transverse line with the anteroposterior centre of the tympanic bulla (2). ORDERED

122. Ventral border of sagittal part of vomer (nasals septum) in ventral view: posteriormost portion projects beyond the posterior border of the palatines and is visible in ventral view (0); completely covered by palatines (1).

123. Basioccipital crest: narrow transversely (0); wide and bulbous (1).

124. Lateral border of basioccipital crest in ventral view: straight (0); concave (1).

125. Orientation of basioccipital crests in ventral view: diverging posteriorly (0); parallel or subparallel (1).

126. Ventromedial corner of paroccipital process in posterior view: located more ventrally than the basioccipital crest (0); level with or more dorsal than the basioccipital crest (1).

127. Posteriormost point of exoccipital in dorsal view: located more anteriorly than posterior edge of occipital condyle (0); level with or posterior to posterior edge of condyle (1).

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