Mammalian Phylogeny

Supplementary Information for the National Science Foundation Proposal:
COLLABORATIVE RESEARCH : A MORPHOLOGICAL DATABASE FOR
THE HIGHER-LEVEL RELATIONSHIPS OF FOSSIL AND RECENT MAMMALS

Principal Investigators and Institutions:

Dr. M. J. Novacek: Curator, Division of Paleontology, American Museum of Natural History Central Park West. at 79th Street, New York, NY 10024.

Dr. G.W. Rougier: Assistant Professor, Department of Anatomical Sciences and Neurobiology, University of Louisville, Louisville 40292 KY.

Dr. J. R. Wible: Curator, Section of mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, PA 15206.

Dr. I. Horovitz: Assistant Curator, Section of Mammalogy, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, CA 90007

Characters:
* Indicates ordered multistate character

6. Posterior nasal expansion width:

0. pronounced, at least twice minimum width of the nasal 1. moderate, less than twice the minimum width 2. uniform width 3. norrows posteriorly

7. Frontonasal suture shape:

0. W-shaped with long medial process of the frontal 1. W-shaped with short medial process ( 1/3 or less the length of the long arm) 2. straight 3. broad v-u shaped 4. narrow v shaped

8. Posterior extent of nasal:

0. posterior to anterior edge of orbit 1. approximately at the level of anterior edge of orbit or in front

9. Nasal foramina:

0. large and multiple 1. small or absent

10. Insicive foramen:

0. approximately the length of the premaxilla, formed by premaxilla-maxilla 1. large, elongated foramen subequal in length to the tooth row 2. large confluent foramen at rostrum's tip 3. absent 4. much smaller than the palatal surface of the premaxilla, subequal to a tooth 5. three small apertures

11. Posterodorsal process of the premaxilla:

0. extends posterodorsally without reaching the frontal 1. extends posteriorly and reaches the frontal 2. weak or absent, suture nearly vertical 3. does not reach the nasals (septomaxilla separates premaxilla from nasal)

12. Number of upper incisors:

0. 5 1. 4 2. 3 3. 2 4. 1 5. none

13. Evergrowing first incisor:

0. absent 1. present

14. Shape of incisor arcade:

0. arched 1. straight 2. U-shaped

15. Frontomaxilla contact:

0. absent 1. narrow, limited by lacrimal and nasals /premaxilla 2. broad, limited by lacrimal and nasals

16. Orbital rim:

0. distinct 1. confluent with rostrum

17. Nasal-facial exposure of premaxilla-maxilla:

0. premaxilla smaller than maxilla 1. premaxilla-maxilla nearly equal in surface

18. Lacrimal:

0. Large facial exposure 1. facial exposure small, confined to rim 2. limited to the orbit 3. absent

19. Number of lacrimal foramina:

0. two 1. one 2. none

20. Position of the lacrimal foramina exposed on the face:

0. on the face, near orbital rim 1. two, one exposed on the orbit, the other on the face 2. on the orbit 3. on the face far from orbital rim 4. on the face on bridge of infraorbital canal

21. Jugal:

0. Present 1. absent

22. Jugal-zygomatic arch:

0. separates maxilla from squamosal 1. reduced, does not separate, jugal on medial side 2. does not separate maxilla, meets squamosal on medial side of arch

23. Maxilla-jugal contact:

0. oblique 1. jugal bifurcates 2. maxilla bifurcates 3. contacts faces ventrally

24. Position of posterior edge of maxilla-jugal contact in lateral view:

0. in front of the last molar 1. at the level or behind the last molar

25. Jugal posterior extent:

0. reaches the glenoid, with a preglenoid process 1. reaches without a preglenoid process 2. does not reach glenoid

26. Infraorbital canal length:

0. long, more than twice the diameter rostral aperture 1. short, less than twice the diameter 2. absent

27. Number of exits of infraorbital canal:

0. two or more 1. single

28. Zygomatic arch incomlete:

0. complete 1. incomplete

29. Septomaxilla:

0. present 1. absent

30. Parietal postorbital process:

0. absent 1. present

31. Multiple row of cusps in molariforms:

0. absent 1. present

32. Cochlear coiling:

0. not coiled (less than 360 degrees) 1. coiled

33. Confluent hypoglossal and jugular foramina:

0. absent 1. present

34. Cochlear aqueduct:

0. absent 1. present

35. Lower jaw coronoid process:

0. present 1. small to absent

36. Anterior lamina:

0. present 1. absent

37. Number of lower incisors:

0. four 1. three 2. two 3. one 4. nore

38. Number of lower molars in the adult:

0. four or more 1. three 2. two 3. one 4. none

39. Carnasial molar:

0. absent 1. present

40. Selenodont dentition:

0. absent 1. present

41. Last premolariform is the only element with deciduous predecesor:

0. absent 1. present

42. Molariform crown simplified with reduced cusps or absence thereof:

0. present 1. absent

43. Inflected mandibular angle:

0. absent 1. present

44. Major axis of the mandibular condyle anteroposteriorly directed:

0. absent 1. present

45. Postorbital ring:

0. absent 1. present

46. Optic foramen:

0. Absent 1. present

47. V3 exits:

0. single 1. multiple

48. Alisphenoid contribution to bulla:

0. week or absent 1. present

49. Petrosal bulla:

0. Absent or very limited contribution 1. present

50. Frontal diploetic system on skull roof:

0. small frontal foramina or absent 1. well developed with large frontal foramina

51. Piriform fenestra:

0. absent or narrow 1. rostral to promontorium 2. rostral and medial to promontorium

52. Echidna pterygoid:

0. present 1. absent

53. Molars:

0. non-tribosphenic 1. tribosphenic, no hypocone on uppers 2. tribosphenic with hypocone. 3. tribosphenic with enlarge cingulum in distolingual position

54. Length of the plagiaulacoid tooth to lower tooth row:

0. long 1. short

55. Prootic canal:

0. long and vertical 1. short and horizontal 2. absent

56. Compound bulla:

0. absent 1. present

57. Last upper pm:

0. simple 1. complex 2. molariform 3. absent

58. Tall trenchant penultimate upper pm:

0. absent 1. present

59. Molariform penultimate upper pm:

0. absent 1. present

60. Palatal vacuities:

0. absent 1. present

61. Lower canine:

0. presence 1. absent

62. Lower canine:

0. tall piercing tooth 1. smaller subequal to incisors and pm

63. Lacrimal jugal- contact:

0. present 1. absent

64. Maxilla excludes jugal from ventral orbital rim.:

0. absent 1. present

65. Lateral flang:

0. present 1. absent

66. Protocone:

0. small or absent 1. large and occluding on talonid

67. Palatal exposure of premaxilla reduced to a narrow strip of bone:

0. 1.

68. Entoglenoid process of the squamosal:

0. Absent or week 1. strong confluent with postglenoid process

69. Ventrally projecting ectotympanic bulla:

0. absent 1. present

70. Fused simphysis:

0. Absent 1. present

71. Character 74:

0. 1.

72. Coracoid and scapular contributions to glenoid fossa:

subequal scapular larger

73. Coronoid process on coracoid*:

absent shorter than glenoid fossa longer than glenoid fossa

74. Coronoid process orientation:

medial anterior lateral

75. Scapular anterior edge recurved:

absent present

76. Scapular spine:

0. absent 1. present

77. Metacromion:

0. absent 1. present

78. Fossae scapulae planes:

same plane different planes

79. Alignment of glenoid fossa major axis and scpular lamina:

aligned not aligned

80. Scapular seccion of glenoid fossa:

wider anteroposteriorly wider dorsoventraly

81. Infraspinous fossa width:

less than half its length more than half its length

82. Spine-scapula connection length:

connection reaches neck connection ends more distally

83. Scapular double spine:

absent present

84. Premetacromion:

absent present

85. Acromion:

absent present

86. Supraspinous fossa anterior border the shortest:

absent present

87. Olecranon fossa/foramen*:

fossa absent fossa present supratroch. for.

88. Humeral fossa on posterior aspect, distal epiphysis, medial trochlea:

absent present

89. Humeral fossa on posterior aspect, distal epiphysis, lateral to trochlea:

absent present

90. Medial epicondyle width (a), relative to mediolateral width of proximal epiphysis (b):

small or absent (a/b £ 0.4) large (0.44 £ a/b )

91. Width of distal epiphysis of humerus excluding medial epicondyle (c), relative to b (see ch. 90):

small (c/b £ 0.85) large (0.9 £ c/b)

92. Entepicondylar foramen:

absent present

93. Supraepicondylar crest:

narrow, non laminar proximal to level of entepicondylar foramen narrow, laminar above level of entepicondylar foramen short, with major axis mediolateral

94. Humeral greater tuberosity:

level or lower than head higher than head

95. Humeral deltopectoral crest*:

absent only proximal extends into distal half

96. Lesser tuberosity:

absent present

97. Proximal extension of humeral condyle and trochlea:

condyle proximal edge at same level or not as proximal as trochlea condyle proximal edge more proximal than trochlea

98. Humeral head:

ovoid (anteropoterior over mediolateral diameters smaller than 1.2) mediolaterally compressed (anteropoterior over mediolateral diameters larger than 1.6)

99. Rim lateral to trochlea:

absent present

100. Lateral area of capitulum is trochlea-shaped:

absent present

101. Lateral extension of capitulum:

absent spiral rod

102. Lip lateral to capitulum flaring laterally:

0. absent 1. present

103. Lip lateral to capitulum:

level lip

104. Proximodistal alignment of proximal and sital humeral epiphyses:

rotated aligned

105. Insertion of teres major on humerus (on medial side of diaphysis):

no trace on bone tuber or ridge groove

106. Ridge for tricipital insertion on lateral side of humeral diaphysis*:

absent present, independent ridge big, lateral edge of deltoid crest process

107. Groove on anterior aspect of lesser tuberosity:

absent present

108. Humeral capitulum shape*:

spherical mediolaterally elongated conical

109. Bicipital groove on head and/or diaphysis*:

absent present on head present on head and shaft

110. Humero-ulnar articulation:

condylar trochlear behind radius

111. Fusion of ulna and radius:

absent present

112. Ulna distal end:

blunt styloid

113. Ulna processes embracing radius:

absent present

114. Ulna, distal postero-medial extension:

absent present

115. Ulna/radius proximal articular facets for humerus proportions:

ulna less than half of radius ulna at least half of radius

116. Ulna facet for humerus position relative to radius

medial posterior

117. Olecranon reduction:

absent present

118. Ulnar distal articular shape:

straight ball-n-socket type saddle hinge

119. Radius-ulna distal end diameter proportions*:

subequal ulna tip smaller ulna shaft smaller ulna splinter

120. Proximal radius process:

absent present

121. Radius proimal. shape:

compressed circular

122. Distal radius:

facet for one bone (scapholunar) facets for two bones (scaphoid and lunar)

123. Lunar proximal facet mediolaterally or anterposteriorly wider:

mediolaterally longer or square anteroposteriorly longer

124. Scaphoid proximal facet shape*:

convex flat or slightly concave hinge (concave)

125. Ulna-lunar contact:

absent present

126. Lunar-unciform contact:

absent present

127. Trapezium-metacarpal contact*:

narrower than unciform-metacapal as wide as unciform-mc widest

128. Trapezium-metacarpal relative positions:

McI radial to Trzm McI parallel to Trzm

129. Mc I-trapezium articulation:

Mc I convex or flat Mc I saddle (ccv apost) Mc I concave-convex

130. Trapezium number of distal heads*:

single two heads present three heads present

131. Centrale:

fused free

132. Centrale position:

prox to trapezoid (trd) lat to trd (trd-scap cont)

133. Capitate size:

less than one fourth the hamate half the size of the hamate or larger

134. Capitate waisted medially:

absent present

135. Capit. narrow prox in dors view

squarish narrowed prox lateral prox tip

136. Proximal phalanges with ventral processes on medial and lateral side:

absent present

137. Hand first digit (phalanges):

absent present

138. Hand fifth digit (phalanges):

absent present

139. Hand fourth digit (phalanges):

absent present

140. Hand second digit (phalanges):

absent present

141. Scaphoid-lunate articulation:

one facet present or fused proximal and distal separate facets

142. Scaphoid-trapezium fusion:

absent present

143. Fingers webbed forming wing:

absent present

144. Obturator foramen subdivision*:

absent incipient complete

145. Anterior inferior iliac origin:

no relief spine pit

146. Position of anterior inferior iliac spine or pit:

next to acetabulum away from acetabulum

147. Ischial-vert.ebral column contact*:

absent posterior to acetabulum acetabulum and posterior region

148. Pubic contact*:

absent vertix side

149. Ischium-postpubic dorsoventral alignment:

pubis more medial pubis aligned or lateral

150. Ilium major transverse axis:

dorsoventral mediolateral

151. Ilium-vertebral contact position:

medial to major face on major face

152. Acetabular notch:

absent present

153. Acetabular notch position:

absent posterior ventral

154. Postobturator notch:

absent present

155. Postobturator foramen:

absent present

156. Ilium anterior and mediolaterally wider than main body of illium:

as wide wider

157. Iliopubic process:

absent present

158. Ischial posterolateral process:

absent present

159. Ischial spine:

absent present

160. Acetabulum: dorsal edge incision (and therefore articular surface is wider anteriorly and posteriorly than in the middle area):

absent present

161. Acetabulum outline:

circular elongate anteroposteriorly

162. Pelvis midline contact*:

absent just ventral ventral midline and posteriorly up to horizontal ischium

163. Acetabulum orientation:

ventrolateral lateral

164. Obturator foramen size:

obturator smaller than acetab. obt. > acetab.

165. Pubic process:

absent present

166. Epipubic bones:

absent present

167. Femur head:

spherical elongated laterally

168. Greater troch two heads:

absent present

169. Greater and lesser trochanter proportions:

different symmetrical

170. Greater trochanter:

lower or equal to head higher than head

171. Lesser trochanter:

just a slight prominence or absent present as a process

172. Lesser trochanter in proximal view:

not directed post directed posteriorly

173. Third trochanter:

absent present

174. Third trochanter position*:

continuous with greater trochanter separate and located in proximal half separate and located in distal half

175. Femur head anteroposterior size:

smaller than greater trochanter larger than greater trochanter

176. Femur medial greater trochanteric fossa:

absent present

177. Femur distal epiphysis width:

wider than deeper deeper than wider or equal

178. Femur head pit for ligament:

surrounded by articular surface not surrounded

179. Femur med lip-lat lip*:

slanted laterally symmetrical slanted medially

180. Patellar facet on femur, width*:

narrower than diaphysis as wide as diaphysis wider than diaphysis

181. Crest liking major and minor trochanter:

absent present

182. Lateral condyle wider:

subequal one third wider

183. Larger trochanter originates lower than head:

0. level with third trochanter 1. higher than third trochanter

184. Posterior trochanteric fossa:

absent present

185. Epiphyses:

absent present

186. Femoral medial condyle more protruding distally:

absent present

187. Femur-fibula articulation:

absent present

188. Fibula posterodistal shape:

simple accessory knob accessory knob protrudes posteriorly

189. Fibula lateral to tibia:

absent present

190. Fibula in tarsus:

absent present

191. Fibula styloid process encases astragalus:

absent present

192. Fibula proximal process*:

absent present at a distance

193. Fibula deep:

narrow as deep as tibia

194. Meniscus fibula-foot:

absent present

195. Tibia distal articular surface:

spiral non-spiral

196. Tibia distal medial (Atim, Szalay, 1994) elongation:

absent present

197. Non helical tibia distally slanted:

straight medially longer

198. Tibia posterior malleolus:

absent present

199. Tibia anterior proximal groove:

absent present

200. Tibia anterior malleolus:

absent present

201. Tibia-fibula fused proximally and distally*:

absent present-distally present proximally and distally

202. Tibia-fibula fusion*:

unfused less than half distally more than half distally

203. Tibia medial malleolus:

absent present

204. Tibia medial malleolus-calcaneum articulation:

absent present

205. Tibia, distal bridge (ossified annular ligament)*:

absent incipient present

206. Tibia length*:

femur longer equal to femur tibia longer

207. Tibia proximal cross section*:

wider than deeper symmetrical mesiolaterally compressed

208. Astragalus medial cavity:

absent present

209. Astragalar fossa for medial malleolus:

absent only proximally only distally both

210. Astragalar lateral process:

absent present

211. Astragalus posterior face sagittal orientation*:

posterolaterla to anteromedial anteroposterior posteromedial to anterolateral

212. Astragalar sustentacular facet location:

on neck on body

213. Astragalar posterior facet:

0. absent 1. concave posteriorly and convex anteriorly

214. Astragalus posterior trochlear shelf:

absent present

215. Astragalar middle and anterior facets separate*:

separate connected together

216. Linkage:

on neck ventral surface only on neck medial surface

217. Astragalus-medial cuneiform-navicular spatial relationship*:

medial cuneiform anerior to navicular medial cuneiform medial to distal region of navicular medial cuneiform medial to all navicular astragalus and medial cuneiform articulate

218. Astragalar neck:

absent present

219. Astragalr neck length:

0 . shorter than trochlea 1. longer than trochlea

220. Facet for navicular shape:

convex saddle very concave with a V-shaped notch trochlea almost flat medially convex, laterally concave

221. Astragalus and navicular ball and socket articulation:

absent present

222. Astrag facet for navicular and trochlea aligned:

absent present

223. Astragalus groove for flexor fibularis muscle:

0. continuous with trochlea 1. distinct from trochlea

224. Astragalar groove for flexor fibularis and/or trochlea extend ventrally:

absent present

225. Astragalus-cuboid articulation:

absent present

226. Astragalus antetrochlea length:

absent present

227. Astragalus trochlea excavated:

absent present

228. Astragalus, angle subtended by trochlea and astragalar-fibular facet angle*:

90 degrees intermediate almost 180 degrees

229. Astragalar canal:

absent present

230. Anterior peroneal tubercle:

absent present

231. Position of anterior calcaneal peroneal tubercle*:

protrudes anteriorly anterior non-protruding at a distance

232. Calcaneus mid-lateral prominence:

absent present

233. Calcaneus-fibula articulation*:

absent present, contact vertical present, fibula dorsal present, calcaneum dislplays an articular process

234. Calcaneus lateral to astragalus:

absent present

235. Calcaneus sustentacular facet position*:

distal anterior to posterio