Supplementary for “A new specimen of Biseridens qilianicus indicates its
phylogenetic position as the most basal anomodont ”
Jun Liu1*, Bruce Rubidge2, Jinling Li1
1
Key Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China (* Author for
correspondence: liujun@ivpp.ac.cn)
2
Bernard Price Institute for Palaeontological Research, University of the Witwatersrand, Private Bag 3,
WITS, Johannesburg, 2050, South Africa
Characters used in the phylogenetic analysis
The number preceding the character definition corresponds to that of the columns
in the data matrix. The characters are cited from different sources: SH: Sidor and
Hopson, 1998; M: Modesto et al., 1999; MR: Modesto and Rybczynski, 2000; R:
Rybczynski, 2000; MO: Modesto et al., 2003; AK: Angielczyk & Kurkin, 2003; A:
Angielczyk, 2004; SR: Sidor and Rubidge, 2006; RSM: Rubidge et al., 2006; Fröbisch,
2007.
When an asterisk following the citation, it denotes that the character definition has
been modified or character state(s) has been added/deleted. Coding of characters is
based on the coding of selected characters in original references, sources listed in the
end of character list, and personal observation.
1. Snout width/height ratio: height greater than width (0), height equal to width
(1), or height less than width (2). (SH: 45)
2. Length of dorsal process of premaxillae: short (0), or long, beyond posterior
margin of external nares (1). (SH: 1*; SR: 2*; RSM: 2*)
3. Premaxilla alveolar margin shape: downturned (0), horizontal or slightly
upturned (1), or greatly upturned (2). (SH: 2*; SR: 3*; RSM: 3*)
4. preorbital region: long, more than half the skull length (0), short (1), or greatly
abbreviated, less than 35% of the skull length (2). (M6*, MR6*, R6*, MO1*,
A6*, F1*)
5. Septomaxilla: is not exposed on lateral surface (0) or escapes to have a short
(1) or long facial exposure (2). (SH: 6; SR: 5*, 6*; RSM: 5)
6. Maxilla contacts prefrontal: absent (0), present (1). (SH: 8; SR: 8; RSM: 7)
7. Supraorbital margin: thin (0) or moderately to greatly thickened (1). (SR: 12;
RSM: 12)
8. Adductor musculature originates on lateral surface of postorbital: absent (0),
present (1), or on both postorbital and postfrontal (2). (SR: 13*, 17*; RSM:
13*)
9. Postorbital ventral portion: sharply tipped (0) or anteroposteriorly expanded
upon the zygoma in lateral aspect (1). (M10, R9, MO12*, A10, F29)
10. Postorbital bar: thin (A-P length less than one-third of height) (0), or thickened
such that A-P length is greater than 40% of its height (1). (RSM: 16)
11. Preparietal: absent (0), present (1). (SH: 48; M19*, MR19*, R18*, MO21,
A20*, SR: 24; RSM: 24*, F31*)
12. Shape of dorsal surface of parietal surrounding parietal foramen: flat (0) or
forms well-defined chimney (1). (SH: 21*; SR: 18*; RSM: 19*)
13. Parietals’ contribution to skull table: shorter anteroposteriorly than broad (0),
or transversely as broad as long (1). (MR16*, R16*, A19*, F32*)
14. Parietal posterolateral process: slender and elongate (0), or short (1). (MR17,
R17, A21, F33)
15. Temporal fenestra: small (0) or expanded posterodorsally (1) so that adductor
musculature origination on squamosal visible in dorsal view. (SH: 14*; SR:
19; RSM: 20)
16. Intertemporal region: wider (0) or narrower (1) than interorbital region. (SH:
18*; SR: 20; RSM: 21)
17. Zygomatic arch: approximately at one level with tooth row, basicranium, and
jaw articulation (0) or displaced dorsally well above those features (1). (SR:
22)
18. Postorbital-squamosal contact: absent ventrally (0), present ventrally (1), or
present ventrally with squamosal extending anteriorly beyond postorbital bar
(2). (M11*, MR10, R10, MO14*, A11, F46*)
19. Anterior extension of anterior ramus of squamosal: stops under temporal
fenestra (0), or beyond the anterior margin of the temporal fenestra (1). (SR:
23*; RSM: 23*)
20. Squamosal zygomatic process: parasagittally deep (0), narrow, rod-like (1), or
transversely expanded (2). (M12, MR11, R11, MO16, A13, F42)
Anomocephalus is coded as ‘0’ following M12.
21. The lateral fossa for the origin of the lateral branch of the M. adductor
mandibulae externus on squamosal: absent (0), present (1). (M14, MR12, R12,
MO18, A14, F38)
Coding of Galeops and Otsheria follows A14.
22. Squamosal posteroventral process: absent (0), or present and extends ventrally
to base of quadrate condyle (1). (M15, MR13, R13, MO17, A15, F40*)
23. Supratemporal: present (0), absent (1). (SH: 22; SR: 25; RSM: 25)
24. Tabular contacts paroccipital process of opisthotic (0) or restricted dorsally
(1). (SH: 54*; SR26*)
25. Length of vomerine process of premaxilla: short (0); long, extending
posteriorly and forming part of the medial margin of the inner choana (1); or
absent in ventral view (2) so that vomer abuts body of premaxilla. (SH: 3*;
SR: 1*; RSM: 1*)
26. Sutural contact of palatine and premaxilla: absent (0) or present (1). (M23,
MR22, R21, MO28, A25, F58)
27. Internal narial shelf: absent (0), narrow and formed by premaxilla, maxilla,
and palatine (1), or well developed and formed primarily by premaxilla (2).
(M22*, MR21*, R20*, MO23, A24, F7)
28. Vomer: paired (0) or unpaired (1). (SH: 25*, 26*; MO25; AK11; SR: 27;
RSM: 26; F50)
29. Vomer internarial part: nearly parallel-sided or slightly expanded backward
(0), widest nearly middle (1), strongly constraining backwards (2). (SH: 23*)
30. Interchoanal portion of vomer where it meets the postchoanal portion: broad
(0), forms median ridge (1). (SH: 23*; RSM: 27)
31. Vomer ventral surface: flat to convex (0), lateral ridges and median trough (1),
vertical ridge (2). (SH: 24*)
32. Choanal and postchoanal portions of vomer meet at similar level on palate (0)
or choanal portion is offset ventrally from postchoanal portion (1). (SR: 28)
33. Lateral margin of the choana formed by the palatine: less than 1/3 (0), over 1/3
(1).
34. Palatal surface of the palatine: without evidence of a keratinized covering (0),
with a rounded, bulbous surface texture that may have had a keratinized
covering (1), relatively smooth and flat, but with fine pitting and texturing
suggestive of a keratinized covering (2). (AK22*, F54*)
Coding of this character follows F54.
35. Palatine widest place: at its approximate midpoint of length (0), widens
anteriorly (1), or width relatively constant for entire length (2). (MR24*, R23*,
A30*, F55)
The coding of Galeops follows F55.
36. Palatine dentition broadly distributed (0), restricted to small area (1), or absent
(2). (SH: 36*; SR: 29; RSM: 28*)
37. Dentition on palatal ramus of pterygoid: present (0), absent (1). (SH: 37; SR:
33)
38. Teeth on transverse flange of pterygoid: present (0), absent (1). (SR: 30; RSM:
29*)
39. Lateral palatal foramen absent (0), present (1). (MO30, AK35*, A33, F57*)
40. Transverse flange of pterygoid projects laterally, free of posterior ramus (0),
projects laterally, bound by posterior ramus (1), ventrally directed (2).
(ME31*, MR28*, R27*, MO31*, A31*, F62*)
41. Position of transverse flange of pterygoid: under posterior half of orbit (0),
under anterior half of orbit (1), or preorbital (2). (SH: 73*; SR31; RSM: 30)
42. Basicranial rami of pterygoids: broadly separated (0), narrowly separated with
median trough formed (1), or broadly contacting anterior to basicranium (2).
(SR: 34; RSM: 32)
43. Basipterygoid articulation located high above primary palate (0), just dorsal to
basicranial ramus of pterygoid (1), or at level basicranial ramus (i.e., suture
visible in ventral view) (2). (SR: 35)
44. Ectopterygoid: extends further posteriorly than palatine (0), or vice versa (1)
in palatal aspect. (M24, MR23, R22, MO29, A26, F60)
45. Ectopterygoid teeth: present (0), absent (1). (SH: 39; SR: 36; RSM: 34)
46. Parasphenoid: excluded from (0) or reaches (1) interpterygoid vacuity. (M32,
MR30, R29, A34, F67)
47. Shape of postparietal: wider than tall (0), approximately square (1), or taller
than wide (2). (SR: 37; RSM: 35)
48. Forward rotation of occiput: none (0), moderate (=vertical) (1), pronounced
(2). (SH: 42; SR: 38; RSM: 36)
49. Paroccipital process orientation: strongly posteroventral and lateral (0),
moderately posteroventral and lateral (1), transverse (2) (SH: 65)
50. Quadrate contact: primarily paroccipital process (0), about equal paroccipital
process and squamosal (1), mostly squamosal (2) (SH: 58*)
51. Stapedial foramen: present (0), absent (1). (SH: 76; SR: 39; RSM: 37)
(This foramen is present in Scylacops, and coded as 0 for all taxa of
Gorgonops here.)
52. Jaw articulation permits strictly orthal closure (0), parasagittal movement (1),
or permits very little parasagittal movement (2). (MR38, R37*, MO13, A41,
F87*)
53. Dentary height in canine versus anterior postcanine regions: nearly equivalent
(0), or shows pronounced difference (1). (SH: 79*; SR: 40; RSM: 38)
54. Dentaries: sutured (0) or fused (1) at symphysis. (M33, MR31, R30, A35,
F76)
I coded Patranomodon as ‘?’ and Ulemica and Galeops as state ‘1’, following A35.
55. Dentary-angular suture runs diagonally across lateral surface of mandible (0)
or posterior margin of dentary deeply incised (1). (SR: 41; RSM: 39)
56. Lateral mandibular fenestra: absent (0), present (1). (SH: 93, 94*; SR: 46)
(M35, MR34, R33, MO38, A37, F75*)
57. Lateral dentary shelf: absent (0), or present (1). (M34, MR32, R31, A36,
F81*)
58. Angular reflected lamina dorsal notch: near articular (0), midway between
articular and dentary (1), close to dentary (2) (SH: 97)
59. Angular with pattern of ridges and fossae on its lateral surface: absent (0),
present (1). (SH: 98*; SR: 42; RSM: 40)
60. Dorsal edge of surangular just posterior to dentary with laterally projecting
ridge: absent (0), or present (1). (SR: 43*; RSM: 41)
61. Foramen between prearticular and angular (sometimes bordered by splenial as
well) on medial surface of lower jaw: absent (0), present (1). (SR: 44; RSM:
42)
62. Articular dorsal process: absent (0), present (1). (SR: 45; RSM: 43)
63. Differentiation of upper tooth row: more than one caniniform teeth (0), one
canine (1), or barely differentiated (2). (SR: 48*)
64. Lower tooth row: prominent canine present (0), barely differentiated (1).
65. Posterior margin teeth arrange in one row (0), two rows (2).
66. Premaxillary teeth number: 5 or 6 (0), 4 (1), 2 or less (2).
67. Upper and lower incisors intermesh: absent (0), present in anterior incisors (1),
present in all incisors (2). (SH: 105*;SR: 49; RSM: 44)
68. Incisor heels: absent (0), present (1) (SH: 106)
69. Upper incisors: much larger (0) or roughly equivalent in size to postcanines
(1). (SR: 50; RSM: 46)
70. Precanine maxillary teeth: present (0), absent (1) (SH: 110)
71. Lower canine: fits into choana (0), or into fossa roofed by premaxilla and
maxilla (1), or passes anterior and external to upper canine (2). (SR: 51; RSM:
47)
72. Upper and lower canines: without heels (0) or small heels present (1). (SR: 52;
RSM: 45)
73. Postcanine diastema on upper jaw: absent (0), present (1).
74. Number of upper postcanines: twelve or greater (0), fewer than 12 (1). (SH:
112; SR: 53; RSM: 48)
75. Fine serrations on marginal teeth present (0), serration absent (1), or coarse
serrations present (2). (M3, MR3, R3, MO9, A3, F17)
Angielczyk, K. D. 2004 Phylogenetic evidence for and implications of a dual origin of
propaliny in anomodont therapsids (Synapsida). Paleobiology 30, 268-296.
Angielczyk, K. D. & Kurkin, A. A. 2003 Phylogenetic analysis of Russian Permian
dicynodonts (Therapsida: Anomodontia): implications for Permian
biostratigraphy and Pangaean biogeography, vol. 139, pp. 157-212.
Fröbisch, J. 2007 The cranial anatomy of Kombuisia frerensis Hotton (Synapsida,
Dicynodontia) and a new phylogeny of anomodont therapsids. Zoological
Journal of the Linnean Society 150, 117-144.
Modesto, S., Rubidge, B. & Welman, J. 1999 The most basal anomodont therapsid
and the primacy of Gondwana in the evolution of the anomodonts.
Proceedings of the Royal Society of London Series B-Biological Sciences 266,
331-337.
Modesto, S. P. & Rybczynski, N. 2000 The amniote faunas of the Russian Permian:
implications for Late Permian terrestrial vertebrate biogeography. In The age
of dinosaurs in Russia and Mongolia (ed. M. J. Benton, M. A. Shishkin, D. M.
Unwin & E. N. Kurochkin), pp. 17-34. Cambridge: Cambridge University
Press.
Modesto, S., Rubidge, B., Visser, I. & Welman, J. 2003 A new basal dicynodont from
the Upper Permian of South Africa. Palaeontology 46, 211-223.
Rubidge, B. S., Sidor, C. A. & Modesto, S. P. 2006 A new burnetiamorph
(Therapsida : Biarmosuchia) from the Middle Permian of South Africa.
Journal of Paleontology 80, 740-749.
Rybczynski, N. 2000 Cranial anatomy and phylogenetic position of Suminia
getmanovi, a basal anomodont (Amniota: Therapsida) from the Late Permian
of Eastern Europe. Zoological Journal of the Linnean Society 130, 329-373.
Sidor, C. A. & Hopson, J. A.. 1998 Ghost Lineages and "Mammalness": Assessing
the Temporal Pattern of Character Acquisition in the Synapsida. Paleobiology
24, 254-273.
Sidor, C. A. & Rubidge, B. S.. 2006 Herpetoskylax hopsoni, a new Biarmosuchian
(Therapsida: Biarmosuchia) from the Beaufort Group of South Africa. In .),
Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and
Reptiles (ed., M. T. Carrano, R. W. Blob, T. J. Gaudin and J. R. Wible), pp.
76-113. Chicago: The University of Chicago Press.
Dimetrodon
Romer, A. S., and L. I. Price. 1940. Review of the pelycosauria. Special papers (Geological
Society of America) 28:1-538.
Tetraceratops
Conrad, J., and C. A. Sidor. 2001. Re-evaluation of Tetraceratops insignis (Synapsida.
Sphenacodontia). Journal of Vertebrate Paleontology 21(3):42A.
Laurin, M., and R. R. Reisz. 1996. The osteology and relationships of Tetraceratops insignis,
the oldest known therapsid. Journal of Vertebrate Paleontology 16(1):95-102.
Biarmosuchus
Chudinov, P. K. 1960. Upper Permian therapsids from the Ezhovo locality.
Paleontologicheskii Zhurnal 1960:81-94.
Ivakhnenko, M. F. 1999. Biarmosuches from the Ocher faunal assemblage of Eastern Europe.
Paleontological Journal 33:289-296.
Herpetoskylax
Sidor, C. A., and B. S. Rubidge. 2006. Herpetoskylax hopsoni, a new Biarmosuchian
(Therapsida: Biarmosuchia) from the Beaufort Group of South Africa. Pp.
76-113. In M. T. Carrano, R. W. Blob, T. J. Gaudin, and J. R. Wible, eds.
Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and
Reptiles. The University of Chicago Press, Chicago.
Titanophoneus PIN 157/1
Syodon PIN 175/2
Orlov, Y. A. 1958. Carnivorous dinocephalians from the fauna of Ishev (Titanosuchia).
Trudy Paleontologischeskogo Instituta, AN SSSR 72:1-114.
Estemmenosuchus PIN 1758/6
Chudinov, P. K. 1960. Upper Permian therapsids from the Ezhovo locality.
Paleontologicheskii Zhurnal 1960:81-94.
Ivakhnenko, M. F. 2000. Estemmenosuchus and primitive theriodonts from the Late
Permian. Paleontological Journal 34 (2): 189-197.
Biseridens IGCAGS V 632, IVPP V 12009
Li, J., and Z. Cheng. 1997. First discovery of eotitanosuchian (Therapsida, Synapsida)
of China. Vertebrata Palasiatica 35(4):268-282.
Anomocephalus
Modesto, S., and B. Rubidge. 2000. A basal anomodont therapsid from the lower
Beaufort Group, Upper Permian of South Africa. Journal of Vertebrate
Paleontology 20:515-521.
Modesto, S., B. Rubidge, and J. Welman. 1999. The most basal anomodont therapsid
and the primacy of Gondwana in the evolution of the anomodonts.
Proceedings of the Royal Society of London Series B-Biological Sciences
266:331-337.
Patranomodon
Rubidge, B. S., and J. A. Hopson. 1996. A primitive anomodont therapsid from the base of
the Beaufort Group (Upper Permian) of South Africa. Zoological Journal of the
Linnean Society 117:115-139.
Otsheria PIN 1758/5
Ivakhnenko, M. F. 2003. Eotherapsids from the East European Placket (Late Permian).
Paleontological Journal 37:S339-S465.
Ulemica PIN 157/5, 1116; 2793/1
Ivakhnenko, M. F. 1996. Primitive Anomodonts, Venyukoviids, from the Late Permian of the
East Europa. Paleontological Journal 30: 575-582.
Suminia
Rybczynski, N. 2000. Cranial anatomy and phylogenetic position of Suminia getmanovi, a
basal anomodont (Amniota : Therapsida) from the Late Permian of Eastern Europe.
Zoological Journal of the Linnean Society 130:329-373.
Galeops
Brinkman, D. B. 1981. The structure and relationships of the dromasaurs (Reptilia:
Therapsida). Breviora 465:1-34.
Eodicynodon
Rubidge, B. S. 1990. The cranial morphology of a new species of the genus Eodicynodon
(Therapsida, Dicynodontia). Navorsinge van die Nasionale Museum, Bleomfontein
7:29-42.
Rubidge, B. S. 1984. The cranial morphology and palaeoenviroment of Eodicynodon barry
(Therapsida: Dicynodontia). Navorsinge van die Nasionale Museum, Bleomfontein
4:325-402.
Gorgonops
Sigogneau, D. 1970. Révision systématique des Gorgonopsiens sud-africains. Cahiers
de Paleontologie: 417.
Data matrix in nexus file
#NEXUS
BEGIN TAXA;
DIMENSIONS NTAX=15;
TAXLABELS
Dimetrodon
Biarmosuchus
Herpetoskylax
Syodon
Titanophoneus
Estemmenosuchus
Biseridens
Anomocephalus
Patranomodon
Otsheria
Ulemica
Suminia
Galeops
Eodicynodon
Gorgonops
;
ENDBLOCK;
BEGIN CHARACTERS;
DIMENSIONS NCHAR=75;
FORMAT DATATYPE=STANDARD MISSING=? GAP=- SYMBOLS="012";
MATRIX
Dimetrodon
0000000000 0000000000 0000000000 0000000000
0000000000 0000000-00 0000010000 00000
Biarmosuchus
0110210000 0100000000 0010000?1? ?100000000
12?01?1111 0010000011 ??1000?011 ?0110
Herpetoskylax
0010210010 110000(01)010 0010?001?? 1110010100
1120101111 ?010100011 0110001011 ?0110
Syodon
1120110201 0110110000 0010200010 1010010?00
1210102111 0000100100 1010002101 11010
Titanophoneus
1120111201 0110110000 0010200010 1010011000
1210102111 0000000100 1?10002101 11010
Estemmenosuchus
11111?1000 01001000?0 0010200110 1010000001
22?0???211 0000100100 1?10002100 20002
Biseridens
1111111100 0100101000 0010000011 1110000000 0???1?0012
1000100210 10101?0?11 10011
Anomocephalus
??11?1??00 ??????1000
00???????? ?????????? ???????0?2 ??0?110??? ??2102001- --011
Patranomodon
11?2?10110 1011101111 0111000011 1110021100
0111100022 100?010211 00210?0??- --011
Otsheria
0111110100 01011?1001 0111011110 ?110021100
01111??012 ???1?????? ??2??100?- ?-01?
Ulemica
0111110100 0101101001 1111011110 1012221101
0111111022 ?201111211 0011110000 -0012
Suminia
1111110100 0101101111 1111011??? ?112221111
0111111022 1101111210 102101000- --012
Galeops
?012010010 0???1?1211 111?1010?1 201002?1?0
021111???? ?10111121? 00210????? -???1
Eodicynodon
10?2?10100 1011101212 1111102011 201112?112 021110?02?
01001112?1 001102???1 -0011
Gorgonops
1010210110 111?100000 ?010100121 1110010000
1210100022 0010000110 ?100000001 00110
;
ENDBLOCK;
BEGIN ASSUMPTIONS;
OPTIONS DEFTYPE=UNORD POLYTCOUNT=MINSTEPS;
ENDBLOCK;