The Phylogeny of Living and Extinct Pangolins (Mammalia, Pholidota) and Associated Taxa: A Morphology Based Analysis

Abstract

The present study was undertaken in order to effect a comprehensive phylogenetic analysis of the order Pholidota, examining seven of the eight currently recognized extant species (absent is Manis culionensis, formerly recognized as a subspecies of Manis javanica) and nearly all the well-known fossil taxa, and employing a wide range of osteological characters from the entire skeleton. In addition, the relationship of pangolins to several putative early Tertiary relatives, including palaeanodonts and the enigmatic “edentate” Eurotamandua joresi, were investigated. The goal of the study was to improve understanding of the systematics and the biogeographic and evolutionary history of the pangolins. A computer-based cladistic analysis of phylogenetic relationships among seven extant species of pangolins, five extinct pangolin species (including all but one of the well-preserved taxa), as well as Eurotamandua and two genera of metacheiromyid palaeanodonts, Palaeanodon and Metacheiromys, was performed based upon 395 osteological characteristics of the skull and postcranial skeleton. Characters were polarized via comparison to the following successive outgroups: the basal feliform carnivoran Nandinia binotata and the hedgehog Erinaceus sp., a eulipotyphlan laursiatherian placental. A revised classification is presented based on the results of the analysis. The results support the monophyly of Pholidota and Palaeanodonta by providing new anatomical characters that can serve to diagnose a pangolin/palaeanodont clade, termed here Pholidotamorpha. Pholidota is defined so as to include all living and fossil pangolins, including all three taxa of middle Eocene “edentates” from the Messel fauna of Germany, among them Eurotamandua joresi. The results do not support the monophyly of the remaining two Messel “edentates” originally placed in the same genus Eomanis, which is restricted to the type species Eomanis waldi. Euromanis, new genus, is named with Eomanis krebsi Storch and Martin, 1994, as the type species, to form a new combination Euromanis krebsi (Storch and Martin, 1994). The analysis strongly supports the monophyly of a crown clade of pangolins diagnosed by many anatomical synapomorphies, the family Manidae. This crown clade is sister to the family Patriomanidae, which includes two Tertiary taxa, Patriomanis americana and Cryptomanis gobiensis, within the superfamily Manoidea. The relationship of the Tertiary European pangolin Necromanis to these two families is unresolved. Within Manidae, the extant species are divided into three well-supported, monophyletic genera, Manis for the Asian pangolins, Smutsia for the African ground pangolins, and Phataginus for the African tree pangolins. The latter two form a monophyletic African assemblage, the subfamily Smutsiinae. The biogeographic implications of this phylogeny are examined. A European origin for Pholidota is strongly indicated. The fossil record of pangolins would seem to support a European origin for the modern forms, with subsequent dispersal into sub-Saharan African and then to southern Asia, and the phylogeny produced in this analysis is consistent with such a scenario.

Appendices

Appendix 1

Specimens examined in connection with the present study

Table 2

* Specimens marked by an asterisk were used to code skull characters in the Gaudin and Wible (1999) study, but were not used to code additional skull or postcranial characters utilized in the present study.

Appendix 2

Listing of characters and character states [beginning with hind limb from distal to proximal, then axial skeleton (except skull) from caudal to cranial, forelimb from proximal to distal, and skull and mandibular characters]. * denotes multistate characters; ** denotes multistate and ordered characters. Abbreviations: UP = ungual phalanges.

• 1. Fissured unguals: absent (0), or present (1).

• *2. Ratio of maximum width vs. maximum height of proximal articular surface on manual and pedal unguals (UP): width and height roughly equivalent, ratio ≥1 (0), manual but not pedal UP compressed mediolaterally, ratio <1 (1), or manual and pedal UP compressed mediolaterally, ratio <1 (2).

• *3. Manual and pedal unguals in dorsal view: waisted, with distinct constriction distal to base (0), manual UP nearly uniform width, constriction rudimentary or absent (1), or manual and pedal UP nearly uniform width, constriction rudimentary or absent (2).

• *4. Shape of manual and pedal subungual process in ventral view: rounded triangular platform, perforated by subungual foramina (0), triangular platform but lacking perforations for subungual foramina, with grooves along either side of subungual process leading to subungual foramina instead (1), narrow subungual process, with grooves on either side leading to subungual foramina (2), or triangular or ovate process, unconstricted by grooves, subungual foramina dorsal to subungual process (3).

• *5. Surface of proximal articulation, manual and pedal intermediate phalanges: unpaired, transversely wide fossae (0), paired, shallow, ovate fossae only slightly longer dorsoventrally than wide, bordered by well-developed lateral ridges (1), or paired fossae deep, elongate dorsoventrally with poorly marked lateral ridges (2).

• 6. Width of distal condyles of manual and pedal intermediate phalanges: narrows dorsally, lateral fossae for tendinous insertion visible in dorsal view (0), or nearly uniform, lateral fossae obscured dorsally by lateral ridges of distal condyles (1).

• 7. Distal condyles of manual and pedal proximal phalanges: with single, transversely elongate, cylindrical condyles (0), or condyles divided into medial and lateral pulleys (1).

• **8. Distal keel on metatarsals and metacarpals: present only as a moderate ridge on ventral half of articulation, distinct fossa on dorsal surface of metacarpal above the condyle (0), ventral ridge on ventral half of articulation hypertrophied, distinct fossa on dorsal surface of metacarpal above the condyle still present (1), or keel extends along entire dorsoventral length of condyle, dorsal fossa above condyle absent (2).

• 9. Width of distal condyles of metatarsals/metacarpals vs. epicondyles: condyles narrower than epicondyles (0), or condyles as wide or wider than epicondyles (1).

• **10. Size of pedal ungual phalanx III (UP III) relative to next largest ungual (UP II or IV): next largest ungual larger than, equal to or only slightly smaller than UP III, greatest length ≥95% of III (0), greatest length of next largest ungual ≥80%, <95% of UP III (1), or UP III enlarged, greatest length of next largest <80% of III (2).

• **11. Size of pedal ungual I: not greatly reduced, ≥70% length of UP III (0), intermediate, ≥40%, <70% length of UP III (1), or greatly reduced, <40% length of UP III (2).

• **12. Shape of proximal phalanges (PP) II-V of pes: elongate, with cylindrical shaft and narrow distal condyles, ratio of maximum width of condyles to maximum length of PP <0.5 (0), intermediate length, ratio of width of condyles to length of PP ≥0.5, <0.9 (1), or wide and short, without cylindrical shaft and with wide distal condyles, ratio of width of condyles to length of PP ≥0.9 (2).

• **13. Relative lengths of proximal (PP) and intermediate (IP) phalanges of pes: proximal phalanges elongate, ratio of length of IP IV to PP IV <1.0 (0), proximal and intermediate phalanges approximately equal in length, ratio of length of IP IV to PP IV ≥ 1.0, <1.2 (1), or proximal phalanges greatly shortened, ratio of length of IP IV to PP IV > 1.2 (2).

• 14. Shape of proximal phalanx I of pes: elongate and slender, ratio of maximum length to maximum proximal depth ≥2.2 (0), or short and deep, ratio of maximum length to maximum proximal depth ≤2.0 (1).

• 15. Shape of proximal margin, proximal phalanx I of pes, in lateral view: convex (0), or concave (1).

• *16. Relative lengths of pedal proximal phalanges: proximal phalanges I–III subequal (0), proximal phalanx I is the longest (1), proximal phalanx II is the longest (2), or proximal phalanx III is the longest (3).

• 17. Relative lengths of metatarsals I and V: I is shorter (0), or V is shorter (1).

• *18. Relative lengths of metatarsals III and IV: III measurably longer, i.e., longer by more than 0.5 mm (0), III and IV subequal (1), or IV measurably longer (2).

• **19. Shape of metatarsals III and IV in dorsal view: ratio of width at midshaft to maximum length ≥20% (0), ratio of width to length <20%, >15% (1), or slender, ratio of width to length ≤15% (2).

• *20. Midshaft width versus depth of metatarsals: all cylindrical or wide and flat, width ≥ depth (0), all but first wide and flat (1), only first and fifth wide and flat, others with depth > width (2), only fifth wide and flat, others with depth > width (3), or all but third wide and flat (4).

• 21. Length of metatarsal III vs. tibia: metatarsal III elongate, ≥30% of tibial length (0), or metatarsal III short, <28% tibial length (1).

• 22. Shape of metatarsal I: straight (0), or curved laterally at its distal end (1).

• 23. Medial depression on ventromedial surface of proximal metatarsal II: present (0), or absent (1).

• 24. Proximal end of metatarsal II: unexpanded transversely (0), or expanded transversely (1).

• *25. Number of contacts between metatarsals II and III, III and IV: one, dorsal only (0), only dorsal contact between II and III, two separate dorsal and ventral contacts between III and IV (1), or two separate contacts present (2).

• 26. Orientation of proximal articular surface of metatarsal III: facet vertical, forms flat contact with lateral cuneiform (0), or articular facet overlaps the dorsal surface of metatarsal III (1).

• *27. Shape of proximal articular surface of metatarsal III in proximal view: roughly dumbbell shape, expanded dorsally and ventrally and constricted in between (0), roughly triangular, narrowing ventrally (1), or quadrangular, without constriction (2).

• 28. Proximal contacts of metatarsal IV: contacts cuboid only (0), or overlapped by lateral cuneiform, contacts both lateral cuneiform and cuboid (1).

• *29. Dorsal surface, proximal end of metatarsal IV: flat (0), with narrow proximodistal groove (1), or markedly concave (2).

• *30. Shape of proximal articular surface, metatarsal IV: quadrangular or triangular (0), triangular or quadrangular with distinct indentation along dorsal margin (1), or irregularly shaped, extended dorsomedially with strongly indented ventromedial margin (2).

• *31. Shape of lateral flange of metatarsal V: elongated distally, straight or curved ventrally at distal end, separated from cuboid facet by groove (0), elongated distally, curved dorsally at distal end, separated from cuboid facet by groove (1), or elongate dorsoventrally, separated from cuboid facet by pit enclosed by dorsal and ventral ridges (2).

• 32. Position of lateral process on metatarsal V: proximal to or at the same level as cuboid articular surface (0), or distal to cuboid articulation (1).

• *33. Shape of cuboid facet of metatarsal V: ovate, transversely compressed, width < depth (0), transversely compressed with width < depth, but expanded ventrally (1), or triangular or quadrangular, width ≅ depth (2).

• 34. Orientation of cuboid facet of metatarsal V: horizontal (0), or tilted proximally and medially (1).

• 35. Articular facets on metatarsal V for cuboid and metatarsal IV: surfaces contiguous but distinct, oriented at different angles and separated by distinct ridge (0), or surfaces continuous, poorly demarcated (1).

• 36. Prehallux: absent (0), or present (1).

• 37. Well-defined articular surface on medial cuneiform for prehallux: absent (0), or present (1).

• 38. Well-defined articular surface on navicular for prehallux: absent (0), or present (1).

• 39. Prehallux shape: strongly concave laterally, convex medially (0), or nearly straight (1).

• *40. Shape of medial cuneiform in medial view: irregular, elongated proximodistally, compressed dorsoventrally (0), triangular, tapered dorsoventrally at its proximal end (1), or quadrangular, dorsoventral depth equivalent distally and proximally (2).

• *41. Shape of distal facet for metatarsal I on medial cuneiform: oval (0), triangular (1), or sigmoid or boomerang-shaped (2).

• **42. Shape and size of navicular and middle cuneiform facets on medial cuneiform: middle cuneiform facet elongated dorsoventrally, larger than navicular facet (0), facets circular, nearly equal in size (1), or navicular facet elongate dorsoventrally, larger than middle cuneiform facet (2).

• 43. Position of middle cuneiform facet and facet for metatarsal II on medial cuneiform: contiguous (0), or separate (1).

• 44. Number of facets for middle cuneiform on the lateral surface of medial cuneiform: one (0), or two (1).

• **45. Dorsal surface of middle cuneiform: elongated dorsoventrally [ratio of width/height <0.9] (0), nearly square, width ≅ proximodistal height [ratio of width/height ≥0.9, ≤1.1] (1), or elongated mediolaterally [ratio of width/height ≥1.2] (2).

• *46. Shape of navicular facet on middle cuneiform: ovate (0), quadrangular (1), elongated dorsoventrally with distinct indentation along the middle of medial edge (2), or L-shaped with ventrolateral indentation and elongate dorsoventrally (ratio of maximum dorsoventral depth to greatest width ≥1.8) (3).

• 47. Shape of articulation between middle and lateral cuneiform: short, occupies less than half of dorsoventral extent of lateral face of middle cuneiform (0), or elongated dorsoventrally, occupies ≥75 % of dorsoventral extent of lateral face of middle cuneiform (1).

• 48. Distinct foramen on dorsal surface of tarsus formed between lateral cuneiform and cuboid: absent (0), or present (1).

• **49. Shape of dorsal surface of lateral cuneiform: elongated proximodistally, ratio of width to height ≤0.8 (0), transverse width ≅ proximodistal height, ratio of width to height ≥1.0, <1.3 (1), or widened transversely, ratio of width to height ≥1.4 (2).

• *50. Shape of navicular facet of lateral cuneiform: triangular or quadrangular (0), butterfly-shaped, expanded transversely on dorsal and ventral ends with concave medial and lateral margins (1), or extended ventrally onto plantar tubercle (2).

• *51. Shape of metatarsal III facet on lateral cuneiform: quadrangular or trapezoidal, with relatively straight medial and lateral edges (0), expanded dorsally and ventrally with sharp medial and lateral indentations (1), or T-shaped (2).

• 52. Plantar process of lateral cuneiform: distinctly separate from distal edge of bone in lateral view (0), or contiguous with distal edge (1).

• 53. Shape of plantar process of cuboid in plantar view: elongated transversely (0), or round (1).

• **54. Orientation of plantar process of cuboid in plantar view: tilts proximomedially (0), horizontal (1), or tilts proximolaterally (2).

• 55. Distal facet of cuboid: deepest point lies at or near medial edge (0), or portion of distal facet that articulates with metatarsal IV with shallow medial extension, deepest part of distal cuboid facet near midline of facet (1).

• 56. Size of calcaneal facet of cuboid: large, maximum length >80% maximum height of cuboid (0), or small, maximum length ≤70% maximum height of cuboid (1).

• *57. Shape of navicular facet of cuboid: facet absent (0), facet elongated dorsoventrally (1), facet elongated proximodistally (2), or facet round or triangular (3).

• 58. Relative size of astragalar and calcaneal facets of cuboid: calcaneal facet clearly larger (0), or astragalar facet as large or larger than calcaneal facet (1).

• 59. Astragalar facet of navicular: facet evenly concave, astragalar head evenly convex (0), or facet concave ventromedially, convex dorsolaterally, astragalar head with large corresponding concavity (1).

• 60. Extent of concavity and convexity on astragalar facet of navicular: concavity extends medial to convexity (0), or concavity restricted to ventral side of convexity (1).

• *61. Distal articular facets of navicular for cuneiforms: medial, middle, and lateral cuneiform facets contiguous, separated by low ridge (0), facets for middle and lateral cuneiform continuous, medial cuneiform facet contiguous, separated by low ridge (1), facets for medial, middle, and lateral cuneiforms continuous, medial/middle cuneiform facet partly separated from lateral cuneiform facet by narrow depression (2), or facets for medial, middle, and lateral cuneiforms continuous, forming a y-shaped surface with facets partly separated by narrow depressions (3).

• **62. Astragalar canal: present (0), rudimentary (1), or absent (2).

• **63. Astragalar trochlea shape in dorsal view: symmetrical, proximodistal depth of medial and lateral trochlea roughly equivalent [ratio of lateral to medial depth <1.2] (0), moderately asymmetrical [ratio of lateral to medial depth ≥1.2, <1.4] (1), or strongly asymmetrical [ratio of lateral to medial depth ≥1.4] (2).

• 64. Proximal edge (=posterior edge) of astragalar trochlea in dorsal view: concave (0), or straight or convex (1).

• 65. Extension of astragalar trochlea onto ventral surface of astragalus: present (0), or absent (1).

• 66. Distinct facet for tendon of m. flexor digitorum fibularis: present (0), or absent (1).

• **67. Proximal extent of ectal facet versus astragalar trochlea: ectal facet distal to or even with proximal edge of trochlea (0), only slightly extended past proximal edge of trochlea [extension beyond trochlea <10% of overall proximodistal length of astragalus] (1), ectal facet extended proximally [extension beyond trochlea ≥10%, <20% length of astragalus] (2), or greatly extended proximally [extension beyond trochlea ≥20% length of astragalus] (3).

• 68. Ratio of maximum dorsoventral depth of astragalus to maximum transverse width: ≤60% (0), or >65% (1).

• 69. Width of astragalar neck vs. maximum width of astragalus: <60% (0), or ≥60% (1).

• **70. Position of astragalar head: medially situated, distance from lateral edge of head to lateral edge of body >40% of overall width of body (0), distance >35%, ≤40% (1), distance >30%, ≤35% (2), distance >25%, ≤30% (3), or displaced laterally, distance from lateral edge of head to lateral edge of body ≤25% of overall width of body (4).

• 71. Shape of astragalar head in distal view: head wide, ratio of maximum transverse width to maximum dorsoventral depth >1.25 (0), or head almost circular, ratio of width to depth <1.25 (1).

• 72. Deep groove for calcaneal-navicular “spring” ligament on ventral margin of astragalar head: absent (0), or present (1).

• 73. Dorsal margin of astragalar head: concave (0), or convex (1).

• **74. Shape of ectal facet of astragalus: very narrow, maximum length more than twice width measured perpendicular to long axis (0), intermediate shape, length less than twice but more than one-and-a half times width (1), or broader, maximum length less than one-and-a-half times width measured perpendicular to long axis (2).

• 75. Orientation of long axis on astragalar ectal facet: proximodistal (0), or proximomedial to distolateral (1).

• *76. Sustentacular facet of astragalus: separate facet centrally located (0), separate facet, displaced distally (1), contiguous with medial navicular and cuboid facet distally (2), or absent (3).

• *77. Shape of sustentacular facet of astragalus: quadrangular, width ≈ length (0), ovate, elongated proximodistally (1), triangular, elongated proximodistally (2), or elongated mediolaterally (3).

• **78. Position of sustentacular facet of astragalus: near lateral edge of astragalar neck (0), in midline of astragalar neck (1), or near medial edge of astragalar neck (2).

• *79. Shape of medial navicular facet on astragalar neck in medial view: narrow dorsally, extended proximally along ventral edge (0), depressed proximodistally along entire length (1), or irregular shape, not markedly depressed (2).

• 80. Astragalus/cuboid contact: absent (0), or present (1).

• 81. Shape of fibular facet of astragalus: oblique, irregularly triangular or teardrop-shaped, without strong vertical moiety distally (0), or crescentic or boomerang-shaped, with concavity facing proximoplantarly or plantarly, or horseshoe-shaped, with concavity facing proximally (1).

• 82. Shape of fibular facet of astragalus and position of central concavity: facet crescentic or boomerang-shaped, with concavity facing proximoplantarly or plantarly (0), or facet horseshoe-shaped, with concavity facing proximally (1).

• **83. Length of calcaneus vs. tibia: calcaneus elongate, ≥45% of tibial length (0), calcaneus of intermediate length, ≥30%, <40% tibial length (1), or calcaneus short, <30% of tibial length (2).

• *84. Tuber calcis shape: nearly cylindrical, dorsoventral depth ≥ transverse width (0), expanded transversely (1), expanded obliquely, greatest depth lies along dorsomedial to ventrolateral axis (2), or expanded obliquely, greatest depth lies along dorsolateral to ventromedial axis (3).

• 85. Dorsal surface of tuber calcis: smoothly rounded (0), or with sharp dorsal ridge (1).

• 86. Length of tuber calcis: ≥50% length of calcaneus (0), or <50% length of calcaneus (1).

• 87. Roughened distal surface of tuber calcis (attachment surface for Achilles’ tendon): vertical or slanted somewhat anteriorly toward plantar surface of calcaneus (0), or enlarged along plantar surface of calcaneus, extends forward more than half calcaneal length (1).

• 88. Orientation of calcaneal cuboid facet: distal and medial (0), or directly distal (1).

• **89. Position of sustentacular facet on calcaneus: situated directly medial to ectal and fibular facets (0), elongated distally, distal portion extends distal to ectal and fibular facets, reaching distal edge of sustentacular flange (1), or situated well distal to ectal and fibular facets, contacting distal margin of calcaneus (2).

• **90. Proximal extent of astragalar and fibular facets of calcaneus: astragalar facet extends further proximally (0), proximal extent of two facets nearly equal (1), or fibular facet extends further proximally (2).

• 91. Position of anterior plantar tubercle vs. cuboid facet of calcaneus: proximal to cuboid facet (0), or even with or distal to cuboid facet (1).

• 92. Shape of cuboid facet of calcaneus: transverse width ≈ dorsoventral depth (0), or elongated transversely (1).

• *93. Peroneal process of calcaneus: absent (0), present as small knob (1), present as low, proximodistally elongated ridge (2), or present as laterally or proximally and laterally elongated flange (3).

• *94. Shape of fibular shaft: compressed anteroposteriorly (0), cylindrical (1), cylindrical distally, compressed mediolaterally in its proximal portion (2), or compressed mediolaterally (3).

• 95. Distinct lateral process on lateral malleolus of fibula: absent (0), or present (1).

• 96. Orientation of tibial facet of distal fibula: medial (0), or anteromedial (1).

• 97. Shape of tibial facet of distal fibula: triangular (0), or elongated anteroposteriorly (1).

• 98. Proximal extent of astragalar and calcaneal facets of fibula in posterior view: nearly equal (0), or astragalar facet extends further proximally (1).

• 99. Proximal half of astragalar facet of fibula: extended ventrally (0), or ventral extension absent (1).

• *100. Development of processes on posterior surface of proximal fibula: small, simple posterior process immediately distal to proximal tibial facet (0), posterior process immediately distal to proximal tibial facet, process marked by elongated posterior groove(s) bounded by lateral ridges (1), posterior process immediately distal to proximal tibial facet, marked by short groove bounded by lateral ridges (2), posterior process well separated from proximal tibial facet, process marked by short groove bounded by lateral ridges (3), posterior process well separated from proximal tibial facet, unmarked by posterior grooves (4), or posterior process absent (5).

• 101. Shape of posterior process on proximal fibula: narrow proximodistally (0), or elongate longitudinally (1).

• *102. Anterolateral eminence on proximal fibula: absent (0), present, extends anterior to tibial facet (1), or present, coincident with tibial facet (2).

• **103. Cnemial crest of tibia: sharp, raised, deeply excavated laterally (0), moderately developed with weak proximal lateral excavation (1), or weak, rounded, lacking lateral excavation (2).

• *104. Development of anterior distal and anterior lateral distal processes of tibia: anterior distal process elongated, narrow, anterior lateral distal process absent (0), anterior distal process present but shallow, wide, anterior lateral process absent or rudimentary (1), strongly reduced anterior distal process, anterior lateral distal process well developed (2), or both processes absent (3).

• 105. Posterior distal tibial process: absent (0), or present (1).

• *106. Grooves for tendons of m. tibialis posterior and m. flexor tibialis on medial malleolus of distal tibia: groove on posterior tibial surface lateral to medial malleolus (0), single groove on posterior edge of medial malleolus (1), or two grooves, with separate posterior groove for m. tibialis posterior on posterior edge of medial malleolus, and anterior groove delimited by distinct anterior and posterior ridges for m. flexor tibialis on medial surface of medial malleolus (2).

• 107. Distal extent of anterior and posterior projections of medial malleolus of distal tibia: distal extent of two processes nearly equal (0), or posterior process extends further distally (1).

• 108. Depth of groove for m. tibialis posterior tendon: shallow (0), or deep, closed over by soft tissue to form tunnel (1).

• **109. Tibial torsion: distal tibial facets rotated clockwise (on L tibia, counterclockwise on R) approximately 45° relative to proximal articulation (0), moderate tibial torsion, distal facets rotated 15–30° (1), or little tibial torsion, distal facets rotated 0–5° (2).

• **110. Dorsoventral depth of distal tibial articulations for medial and lateral trochleae of astragalus: medial facet depth > lateral facet (0), facets of approximately equal depth (1), or lateral facet depth > medial facet (2).

• 111. Distal tibial articulation for medial trochlea of astragalus overlaps anterior face of medial malleolus’ posterior extension: absent (0), or present (1).

• 112. Distal tibial articulation for fibula: not visible in distal view (0), or visible (1).

• **113. Maximum width vs. anteroposterior depth of distal tibia: ratio of width to depth <1.5 (0), ratio of width to depth ≥1.5, <2 (1) or distal tibia compressed, ratio of width to depth ≥2 (2).

• **114. Relative size of lateral and medial condyles of tibia in proximal view: medial condyle larger than lateral condyle (0), subequal (1), or lateral condyle larger than medial condyle (2).

• **115. Shape of tibial lateral condyle in proximal view: elongated transversely (0), transverse width and anteroposterior depth roughly equivalent (1), or elongated anteroposteriorly (2).

• *116. Surface contour of tibial lateral condyle: concavo-convex (0), concave (1), convex (2), or flat (3).

• 117. Pit for attachment of meniscal ligament anterior to medial condyle of tibia: absent (0), or present (1).

• 118. Proximal fibular facet of tibia: not visible in anterior view (0), or visible (1).

• 119. Facet for tibial sesamoid (cyamelle) on proximal fibula: absent (0), or present (1).

• 120. Anterior surface of patella: rugose (0), or with fine longitudinal striations (1).

• 121. Articulations on posterior surface of patella: distinct medial and lateral facets separated by a sharp ridge (0), or medial and lateral facets confluent (1).

• 122. Surface of confluent medial and lateral patellar articulations: convex transversely (0), or nearly flat (1).

• 123. Shape of femoral shaft: slender, minimum width <11% of maximum length (0), or broadened transversely, minimum width >13% maximum length (1).

• **124. Angle between femoral head and shaft: <130° (0), 130–135° (1), or >135° (2).

• **125. Shape of greater trochanter of femur in proximal view: anteroposterior depth > transverse width (0), L-shaped, elongated both anteroposteriorly and transversely (1), or anteroposterior depth ≤ transverse width (2).

• 126. Orientation of patellar groove of femur in anterior view: directed toward greater trochanter, medial edge inclined proximally and laterally (0), or directed toward head, medial edge inclined proximally and medially (1).

• 127. Surface contour of patellar groove of femur in distal view: concave (0), or flat (1).

• **128. Distance between proximal end of femur and third trochanter: <50% of maximum length of femur (0), ≥50, <60% of maximum length (1), ≥60, <70 % of maximum length (2), or ≥70 % of maximum length (3).

• 129. Distance between femoral head (measured from proximal-most point) and tip of lesser trochanter: lesser trochanter close to head, distance <25% maximum length of femur (0), or lesser trochanter further separated from head, distance ≥25% maximum length (1).

• **130. Orientation of lesser trochanter: directed posteriorly, visible posterior to head in proximal view (0), directed posteromedially, partially obscured by head but visible posteriorly in proximal view (1), or directed medially, largely obscured by head but visible medially in proximal view (2).

• 131. Fovea capitis: present (0), or absent (1).

• *132. Trochanteric fossa and intertrochanteric ridge: well developed (0), rudimentary or absent (1), or trochanteric fossa large but intertrochanteric ridge rudimentary or absent (2).

• **133. Maximum depth vs. transverse width of distal femur in distal view: width almost equal to depth, ratio = 1 (0), width greater than depth, ratio 1.20–1.29 (1), or width greatly exceeds depth, ratio ≥1.30 (2).

• 134. Maximum depth of medial vs. lateral condyle of femur in distal view: almost equal, ratio of medial depth to lateral <1.20 (0), or medial condyle deeper, ratio ≥1.20 (1).

• 135. Maximum width of medial vs. lateral condyle of femur in distal view: medial condyle wider, ratio of medial width to lateral ≥1.20 (0), or almost equal, ratio >0.9, <1.20 (1).

• 136. Sacroiliac attachment: unfused (0), or fused (1)

• **137. Extent of sacroiliac junction: ends anterior to acetabulum (0), extends to midpoint of acetabulum (1), or extends past midpoint of acetabulum, sacral vertebrae contact anterior ischium (2).

• *138. Number of sacral vertebrae: 2, both attached to ilium (0), 3, 2 attached to ilium (1), 3, all attached to ilium (2), 4, 3 attached to ilium (3), or 4, 2 attached to ilium (4).

• *139. Metapophyses and neural spines of sacral vertebrae: unfused (0), neural spines of two or more sacral vertebrae fused, but metapophyses unfused (1), metapophyses of two or more sacral vertebrae fused, but neural spines unfused (2), or metapophyses and neural spines fused (3).

• 140. Dorsal spinal nerve foramina of sacral vertebrae: face dorsally, well separated laterally from base of metapophyses (0), or face dorsolaterally, situated immediately underneath metapophyses (1)

• 141. Process for sacrospinous ligament at posteroventral base of transverse process of last sacral vertebra: absent (0), or present (1).

• *142. Shape of transverse process of last sacral vertebra: wing-like, expanded anteroposteriorly but thin dorsoventrally (0), unexpanded, rod-like (1), or expanded anteroposteriorly and thick dorsoventrally (2).

• 143. Lateral extent of transverse process of last sacral vertebra in dorsal view: does not extend lateral to body of ischium or acetabulum (0), or extends lateral to ischium (1).

• 144. Metapophyses of sacral vertebrae: short, less than half neural spine height (0), or elongated, >2/3 neural spine height (1).

• **145. Angle between sacral vertebrae and ilium: ≤10° (0), >20°, ≤30° (1), or >35° (2).

• *146. Gluteal fossa on ilium: large, with prominent lateral iliac crest, medial dorsal flange of ilium and caudal dorsal iliac spine (0), weak, iliac crest rounded but still well developed, small dorsal flange and free caudal dorsal iliac spine (1), poorly demarcated, iliac crest rounded, weak, dorsal flange absent, caudal dorsal iliac spine incorporated in sacroiliac junction (2), or poorly demarcated, iliac crest rudimentary but small dorsal flange and caudal dorsal iliac spine present (3).

• *147. Position of femoral spine (= tuberosity for m. rectus femoris): on acetabular eminence, directly dorsal to iliopubic eminence (0), on lateral iliac crest, anterior to acetabular and iliopubic eminence (1), or absent (2).

• 148. Shape of lunate surface of acetabulum: U-shaped, broadly open (0), or C-shaped, nearly a closed loop (1).

• **149. Orientation of acetabular fossa: fossa opens ventrally (0), posteroventrally (1), or posteriorly (2).

• 150. Size of obturator foramen: large, maximum diameter of acetabulum <75 % that of obturator foramen (0), or small, maximum diameter of acetabulum ≥75 % that of obturator foramen (1).

• 151. Orientation of ischial tuberosities: not flared laterally, or only weakly so (0), or flared laterally (1).

• 152. Development of ischial spine: small (0), or prominent (1).

• 153. Position of ischial spine: separated anteriorly from ischial tuberosity, dorsal to anterior portion of obturator foramen (0), or close to ischial tuberosity, dorsal to posterior portion of obturator foramen (1).

• 154. Position of dorsal edge of ischium: at same level as transverse processes of sacral and anterior caudal vertebrae (0), or ventral to transverse processes of sacral and anterior caudal vertebrae (1).

• *155. Shape of pubis and position of attachment to ilium: rod-like, elongated, attached to ilium under anterior edge of acetabulum (0), slightly compressed mediolaterally, attached to ilium beneath midpoint of acetabulum (1), short, flat, attached to ilium under posterior edge of acetabulum (2), or elongated, flat, attached to ilium under anterior edge of acetabulum (3).

• 156. Anterior edge of pubic symphysis in ventral view: forms narrow V (0), or forms broad U (1).

• **157. Number of thoracic vertebrae: 11 (0), 12 (1), 13 (2), 14 (3), 15 (4), or 16 (5).

• **158. Number of lumbar vertebrae: 7 (0), 6 (1), or 5 (2).

• **159. Number of caudal vertebrae: more than 30 (0), 20–30 (1), or less than 15 (2).

• **160. Position of diaphragmatic vertebra: T9 (0), T10 (1), T11 and T12 (1), T12 (2), T12 and T13 (3), or T13 (4).

• *161. Transverse process of last lumbar vertebra: rodlike, reduced in lateral and anteroposterior extent (0), reduced in anteroposterior extent versus that of penultimate lumbar (1), reduced in lateral extent versus that of penultimate lumbar (2), or unreduced (3).

• 162. Transverse processes of anterior lumbar vertebrae: directed anterolaterally (0), directed laterally (1), or directed posterolaterally (2).

• 163. Attachment of pedicel to lumbar centra: ventrolateral to dorsal edge of centrum, spinal nerves form grooves on posterodorsolateral surface of centrum (0), or pedicels attach to dorsal edge of centrum, spinal nerve foramina dorsal to centrum (1).

• 164. Shape of centrum of third lumbar vertebra: short anteroposteriorly, length ≤1.5 times maximum depth (0), or elongated anteroposteriorly, length >1.5 times maximum depth (1).

• **165. Neural spine of third lumbar vertebra: elongated dorsoventrally, height >1.5 times anteroposterior length (0), low, height 1.0–1.5 times anteroposterior length (1), or elongated anteroposteriorly, height <0.7 times anteroposterior length (2).

• 166. Surface contour of zygapophyses of lumbar vertebrae: flat or slightly concave (0), or embracing (1).

• 167. Relative heights of neural spines of anterior thoracic vertebrae: elongated, much longer than in more posterior thoracics (0), or not dramatically elongated (1).

• **168. Anteriormost thoracic vertebra in which the diapophysis is completely separated posteroventrally from the metapophysis: T10 (0), T11 (1), T12 (2), T13 (3), or T14 (4).

• **169. Anteriormost thoracic vertebra in which the capitular facet is entirely confined to a single centrum: T10 (0), T11 (1), T12 (2), T13 (3), T14 (4), or T15 (5).

• 170. Surface contour of anterior and posterior zygapophyses of cervical vertebrae: flat (0), or anterior concave, posterior convex (1).

• 171. Cervical anterior and posterior zygapophyses extend far anterior and posterior to centrum: absent (0), or present (1).

• 172. Heights of neural spines in cervical vertebrae 3-7: C3>C4>C5<C6<C7 (0), or C3≤C4≤C5<C6<C7 (1).

• **173. Inclination of neural spine of C3: vertical (0), weakly inclined posteriorly (1), or strongly inclined posteriorly (2).

• 174. Neural spine of C7: vertical (0), or inclined posteriorly (1).

• 175. Transverse process of C7: rectangular (0), or flared distally (1).

• *176. Posterior edge of lamina of C4-6 in dorsal view: straight (0), concave (1), or irregular (2).

• **177. Posterior extension of neural spine of axis: absent, posterior surface of neural spine with two oval concavities for attachment of nuchal ligament (0), deep, blunt extension over anterior portion of C3 (1), or elongated, narrow extension to midpoint of C3 or beyond (2).

• **178. Anterior extension of neural spine of axis: absent (0), present but small, little overlap of posterior edge of atlas (1), present, strongly overlaps posterior portion of atlas with surface for ligamentous attachment to arch of atlas (2), or present, elongated and narrow with diamond-shaped anterior expansion for ligamentous attachment to arch of atlas (3).

• 179. Ventral surface of transverse process of axis: flat or convex (0), or with longitudinal concavity (1).

• 180. Ventral surface of axial centrum: with midline crest or eminence and lateral excavations (0), or surface relatively smooth, eminence and crest rudimentary or absent (1).

• 181. Ratio of axial centrum maximum anteroposterior length (including dens) to width (measured immediately behind anterior articular surface): ≥1.0 (0), or <0.85 (1).

• *182. Surface contour of anterior articular surface of axis: flat or convex transversely (0), convex medially, concave laterally (1), or concave transversely (2).

• 183. Shape of anterior articular surface of axis: elongated transversely, ratio of width to maximum dorsoventral depth >1.30 (0), or ratio of width to depth <1.25 (1).

• 184. Axial anterior articular surface and articular facet of dens: contiguous (0), or separate (1).

• 185. Transverse foramen of axis in anterior view: not visible, obscured by anterior articular surface (0), or visible (1).

• 186. Transverse process of atlas: broad dorsoventrally (0), or shallow dorsoventrally (1).

• 187. Anterodorsomedial groove connecting transverse canal of atlas with lateral vertebral foramen: open (0), or enclosed by contact between anteromedial edge of transverse process and arch of atlas (1).

• 188. Distinct depressions for origin of mm. rectus capitis anterior and lateral to neural spine of atlas: absent (0), or present (1).

• **189. Maximum width vs. maximum anteroposterior length of atlas (excluding transverse processes): narrow transversely, ratio <1.0 (0), ratio 1.5–2.0 (1), or widened transversely, ratio >2.0 (2).

• 190. Separation of anterior zygapophyses of atlas: widely separate, distance between medial edge of two facets ≥40% of maximum width of atlas (0), or narrowly separated, distance between medial edge of two facets <40% of maximum width of atlas (1).

• 191. Relationship of sternal ribs and sternum: ventral surface of sternum flat, sternal ribs articulate with lateral surface of sternebrae (0), or ventral surface of sternebrae with large, anteroventral processes, sternal ribs overlap posteroventral surface of sternebrae, contact or nearly contact one another in midline (1).

• 192. Shape of manubrium in cross-section: cylindrical (0), or flattened dorsoventrally (1).

• *193. Xiphisternum: short (0), elongated (1), or elongated and bifurcate (2).

• *194. Cartilaginous extension of xiphisternum: short, length much less than ossified portion of xiphisternum, shovel shaped at distal end (0), elongated, length much greater than ossified portion of xiphisternum, shovel shaped at distal end with central perforation (1), or greatly elongated, extending posteriorly to pelvis, then curling dorsally toward vertebral column at its distal end (2). (see Grassé 1955; Kingdon 1971.)

• **195. Number of true ribs: 6 (0), 7 (1), 8 (2), or 9 (3).

• 196. Shape of first rib: shaft expanded mediolaterally (0), or unexpanded, or only slightly expanded at distal end (1).

• 197. Clavicle: present (0), or absent (1).

• 198. Shape of ventral end of scapular spine: elongated acromion process present (0), or acromion rudimentary (1).

• **199. Shape of acromion: blunt, without any indication of bifurcation (0), weakly bifurcated (1), or strongly bifurcated with well-developed metacromion process (2).

• **200. Coracoid process of scapula: present, robust (0), rudimentary (1), or absent (2).

• **201. Distance from ventral corner of anterior scapular edge to anterior edge of glenoid: close, distance <15% of maximum length of scapula [measured from ventral edge of glenoid to dorsal edge of scapula] (0), distance ≥15%, <20% (1), distance ≥25%, <35% (2), or widely separated, distance ≥55 % of maximum length of scapula (3).

• **202. Secondary scapular spine: sharp crest (0), rounded crest (1), or absent or rudimentary (2).

• 203. Teres minor fossa on posteroventral edge of scapula: present (0), or absent (1).

• 204. Height of scapular spine in distal view: elongated, height roughly equivalent to mediolateral width of glenoid (0), or reduced in height, <85% of mediolateral width of glenoid (1).

• 205. Orientation of long axis of humeral head in posterior view: oriented proximodistally, or somewhat distolateral (0), or oriented distomedially (1).

• **206. Orientation of deltopectoral crest of humerus: canted medially at its distal end (0), extends straight down shaft (1), or canted laterally at its distal end (2).

• **207. Length of deltopectoral crest: extends >75% of the length of the humerus (0), extends >65, ≤75% of the length of the humerus (1), extends >55, ≤65% of the length of the humerus (2), or extends ≤55% of the length of the humerus (3).

• 208. Pulley for m. biceps brachii at distal termination of deltopectoral crest: present (0), or absent (1).

• 209. Deep anterior groove between greater tubercle and lesser tubercle of humerus for head of m. biceps brachii: present (0), or groove shallow or absent altogether (1).

• 210. Large distinct deltoid crest extends across lateral surface of humerus: present (0), or absent (1).

• 211. Distinct fossa for mm. infraspinatus and teres minor on lateral surface of humerus: absent (0), or present (1).

• **212. Width across humeral epicondyles: relatively narrow, <40% of length of humerus (0), intermediate width, 40–50% of length of humerus (1), or wide, >50% of length of humerus (2).

• **213. Length of entepicondyle: short, <30% of epicondylar width of humerus (0), intermediate length, 30–40% of epicondylar width of humerus (1), or elongated, >40% of epicondylar width of humerus (2).

• 214. Position of proximal entepicondylar foramen: visible in anterior view (0), or visible in posterior view (1).

• *215. Entepicondylar notch: absent (0), weak (1), or present (2).

• **216. Size and proximal extent of supinator crest: greatly enlarged, with free-standing proximal extension reaching to middle region of humeral shaft (0), well developed but lacking free-standing proximal extension, extends proximal to proximal opening of entepicondylar foramen and extends far lateral to humeral shaft (1), moderately developed, extends proximal to proximal opening of entepicondylar foramen but weakly flared laterally (2), or reduced, ends at level of proximal opening of entepicondylar foramen (3).

• 217. Form of supinator crest: flared laterally at its distal end, with well-developed anterior concavity (0), or extends nearly straight distally, anterior concavity poorly marked or absent (1).

• 218. Posterior surface of distal humerus between supinator crest and posterior edge of entepicondyle/entepicondylar foramen, proximal to olecranon fossa: flat (0), or concave (1).

• 219. Distal edge of trochlea of humerus in anterior view: straight or slightly concave (0), or convex (1).

• 220. Medial lappet of trochlea of humerus extending underneath entepicondyle in distal view: present (0), or absent (1).

• 221. Medial extent of radial fossa of humerus: extends over lateral half of trochlea (0), or situated directly above capitulum, does not extend medially over trochlea (1).

• 222. Size of olecranon fossa of humerus: large, wider transversely than trochlea in posterior view (0), or small, narrower transversely than trochlea (1).

• 223. Relative proximal extent of greater and lesser tubercle of humerus: greater tubercle extends proximal to lesser (0), or proximal extent of greater and lesser tubercle roughly equivalent (1).

• 224. Position of proximal portion of lesser tubercle of humerus: extends anterolaterally, overlaps head in anterior view (0), or does not extend anterolaterally, remains medial to head in anterior view (1).

• 225. Bicipital groove of humerus continuous with well-developed fossa anteromedial to greater tubercle: present (0), or absent (1).

• 226. Orientation of greater tubercle of humerus relative to head in proximal view: divergent from head posterolaterally (0), or approximately parallel to lateral surface of head (1).

• **227. Length of olecranon process of ulna: short, <20% of maximum ulnar length (0), intermediate length, 20–30% of maximum ulnar length (1), or elongated, >30% of overall ulnar length (2).

• 228. Olecranon process of ulna inflected medially in anterior view: absent (0), or present (1).

• 229. Olecranon process of ulna inflected ventrally in medial view: absent (0), or present (1).

• **230. Maximum width of radiohumeral articular surface of ulna: wide, ≥20% of maximum ulnar length (0), intermediate, ≥15%, <20% of maximum ulnar length (1), or narrow, ≤10% of maximum ulnar length (2).

• **231. Width of anconeal process of ulna: narrow, <10% of maximum ulnar length (0), 10–15% of maximum ulnar length (1), or wide, >15% of maximum ulnar length (2).

• 232. Distance between coronoid process and ulnar tubercle of ulna: close, ≤10% of maximum ulnar length (0), or biceps tubercle displaced distally, distance >10 % of maximum ulnar length (1).

• *233. Relative anterior extent of anconeal and coronoid processes of ulna: anconeal process extends further anteriorly (0), coronoid process extends further anteriorly (1), or equal in anterior extent (2).

• 234. Shape of interosseous border of ulna: sharp (0), or rounded, poorly marked (1).

• **235. Medial concavity on distal ulnar shaft: well marked (0), weakly present (1), or absent (2).

• 236. Depth of distal ulnar shaft: deep, ≥10% of maximum ulnar length (0), or narrow, <10% of maximum ulnar length (1).

• 237. Shape of styloid articulation of ulna in distal view: maximum transverse width ≥ anteroposterior depth (0), or compressed mediolaterally, width < depth (1).

• **238. Ratio of ulnar length to humeral length: ≤90% (0), >90, ≤100% (1), >100, ≤120% (2), or >120% (3).

• **239. Ratio of radial length to humeral length: <75% (0), >75, ≤85% (1), or >95% (2).

• **240. Maximum depth of radial shaft: <15% of maximum radial length (0), ≥15, <20% of maximum radial length (1), ≥20, <25% of maximum radial length (2), or ≥25% of maximum radial length (3).

• *241. Radial (=bicipital) tuberosity: absent (0), present, weakly developed (1), strong, angular process (2), or present as a well-developed tubercle (3).

• 242. Size of trochlear facet on radial head: large, ≥1/3 the overall width of the head (0), or small, <1/3 the overall width of the head (1).

• 243. Ulnar facet and sesamoid facet on radial head: contiguous (0), or separate (1).

• **244. Sesamoid facet on radial head: absent (0), small, not visible in proximal view (1), or large, visible in proximal view (2).

• 245. Condylar facet of radial head: concave medially with flat lateral portion (0), or evenly concave (1).

• *246. Styloid and pseudostyloid processes and dorsal tuberosity of distal radius: styloid process rudimentary, pseudostyloid process prominent, dorsal tuberosity weak (0), styloid process, pseudostyloid process and dorsal tuberosity all prominent (1), styloid process and dorsal tuberosity prominent, pseudostyloid process rudimentary or absent (2), dorsal tuberosity prominent, much larger than styloid process, pseudostyloid process weakly developed (3), dorsal tuberosity prominent, styloid process and pseudostyloid process weakly developed (4), or styloid process prominent, pseudostyloid process and dorsal tuberosity weakly developed or absent (5).

• 247. Orientation of radial styloid process: distal (0), or distolateral (1).

• **248. Orientation of distal radial articular surface: long axis transverse (0), long axis runs dorsomedial to ventrolateral (1), or long axis runs dorsoventral, or nearly so (2).

• 249. Fused scaphoid and lunate bone: absent (0), or present (1).

• 250. Contact between scapholunar and hamate: present (0), or absent (1).

• 251. Proximodistal depth of palmar tuberosity on scapholunar: palmar tuberosity narrow, depth <30% of maximum width of scapholunar (0), or palmar tuberosity deep, depth >35% of maximum width of scapholunar (1).

• 252. Shape of palmar margin of radial articular surface of scapholunar: nearly straight (0), or strongly indented, concave in palmar direction (1).

• 253. Triquetral articular surface on scapholunar: absent (0), or present (1).

• 254. Connection between trapezoid and capitular articular surfaces on scapholunar: capitular divided from trapezoid facet by sharp ridge (0), or trapezoid and capitular facets continuous (1).

• 255. Shape of ulnar articular facet of triquetrum: compressed dorsoventrally, transverse width much greater than dorsoventral depth (0), or width ≤ depth (1).

• 256. Ulnar/pisiform articular facets vs. scapholunar facet on triquetrum: facets separate (0), or facets contiguous (1).

• *257. Surface contour of scapholunar facet on trapezium: convex (0), concave (1), or complex, concavo-convex (2).

• 258. Extent of scapholunar facet on trapezium: restricted to medial portion of proximal surface (0), or covers entire proximal surface of bone (1).

• 259. Surface contour of scapholunar facet on trapezoid: concave (0), or convex (1).

• **260. Orientation of scapholunar facet on trapezoid in dorsal view: facet tilted to face proximally and laterally (0), facet horizontal (1), or tilted to face proximally and medially (2).

• **261. Orientation of facet for trapezium on trapezoid: distally and medially (0), medially (1), or proximally and medially (2).

• **262. Width of proximal articulation on capitate: narrow, width <50% of maximum dorsoventral depth of capitate (0), ≥50%, <60% (1), ≥60%, <70% (2), ≥70%, <80% (3), or very wide, ≥85% (4).

• *263. Shape of proximal articulation on capitate: ovate or rectangular, without lateral indentation (0), roughly triangular, with slight lateral indentation (1), or L-shaped, with abrupt, step-like lateral indentation between dorsal portion of facet and its palmar extension (2).

• *264. Relationship of capitate and trapezoid: trapezoid overlaps capitate proximally, trapezoid facet of capitate faces proximally and medially (0), capitate overlaps trapezoid proximally, trapezoid facet of capitate faces distally and medially (1), or capitate and trapezoid abut along a vertical surface (2).

• *265. Shape of distal articulation of capitate: dumbbell shape, with indentations along the medial and lateral margins (0), roughly triangular, narrower ventrally than dorsally (1), bilobate, with an indentation along the medial margin only (2), or rectangular (3).

• 266. Position of hamate facet of capitate: near dorsal edge of bone, dorsal to dorsoventral midline (0), or displaced ventrally, lies at or ventral to dorsoventral midline (1).

• *267. Unciform process of hamate: absent (0), directed distally and ventrally (1), or directed distally and ventrally at its base, with a distinct proximal and ventral extension (2).

• 268. Dorsal surface of hamate: narrow proximodistally, proximodistal length < transverse width (0), or narrow transversely, length ≥ width (1).

• 269. Facet for metacarpal IV on hamate: narrow transversely (width < dorsoventral depth) (0), or wide transversely (1).

• 270. Metacarpal III length vs. humerus: III elongated, ≈ 30% of humerus length (0), or short, <25% of humerus length (1).

• 271. Shortest metacarpal: V (0), or I (1).

• 272. Longest metacarpal: IV (0), or III (1).

• **273. Relative length of metacarpal II and IV: II < IV (0), II = IV (1), or II > IV (2).

• **274. Width vs. length of metacarpal IV: narrow transversely, minimum width <20% of maximum length (0), minimum width ≥20, <30% of maximum length (1), minimum width ≥30, <45% of maximum length (2), or expanded transversely, minimum width ≥45% of maximum length (3).

• *275. Cross-sectional shape of shaft of metacarpals I-V: roughly cylindrical, width ≥ depth (0), dorsoventrally elliptical, depth > width (1), or depth > width on all but metacarpal III, in which width ≥ depth (2).

• *276. Articulation between distal metacarpals and proximal phalanges: distal condyles and keels of metacarpals convex, joint mobile (0), all but metacarpal III with mobile joints, metacarpal III with nearly vertical keel and ventral stop, joint immobile (1), or joint surface flat or concavoconvex on metacarpals I-IV, joints immobile (2).

• *277. Shape of metacarpal I in medial view: with elongated cylindrical shaft, may be slightly expanded proximally and distally (0), dumbbell shaped, with broadly expanded proximal and distal ends joined by short shaft (1), or expanded distally only, with shaft narrowing towards proximal end (2).

• 278. Shape of proximal articular surface of metacarpal I in proximal view: triangular, transverse width ≈ dorsoventral depth (0), or compressed transversely (1).

• 279. Distal keel on condyle of metacarpal I: present (0), or absent (1).

• 280. Concavity on dorsal surface of metacarpal II, proximal end: absent (0), or present (1).

• 281. Facet for trapezium on metacarpal II: restricted to dorsal half of medial surface (0), or extends almost to ventral extremity of medial surface (1).

• *282. Articulation between metacarpal II and III: articular surfaces curved, concave on metacarpal II, convex on metacarpal III (0), articular surfaces flat (1) or facet convex on metacarpal II, concave on metacarpal III (2).

• 283. Palmar extent of capitular facet on metacarpal III: palmar tuberosity well developed, little overlap of capitular facet on shaft (0), or facet overlaps shaft proximally, palmar tuberosity weakly developed (1).

• 284. Shape of capitular facet of metacarpal III in medial view: evenly convex (0), or sigmoid, extended to form dorsal shelf (1).

• 285. Dorsal surface of metacarpal III, proximal end: smoothly convex (0), or with sharp midline crest terminating in prominent tubercle proximally (1).

• 286. Proximal and lateral extension of shaft of metacarpal III overlapping metacarpal IV proximally, lateral to capitate facet: present (0), or absent (1).

• 287. Hamate facet of metacarpal III: elongated dorsoventrally (0), or restricted to dorsal half of proximomedial surface (1).

• *288. Extensor tubercle on dorsal surface of metacarpals: present on metacarpals III and IV (0), present on metacarpals II and III (1), or absent (2).

• *289. Dorsal surface of metacarpal IV extended laterally and medially for articulations with metacarpals III and V: absent (0), T-shaped proximal end, extensions present medially and laterally (1), or extension present laterally but not medially (2).

• *290. Surface contour of proximal articulation of metacarpal IV: evenly convex transversely and anteroposteriorly (0), mostly convex but with strong concave pit (1), or concave transversely, convex anteroposteriorly (2).

• *291. Shape of proximal articulation of metacarpal IV: rectangular, long axis oriented dorsoventrally (0), triangular with base facing dorsally (1), or semicircular with flat edge facing dorsally (2).

• 292. Contour of articular surface for metacarpal III on metacarpal IV: flat dorsoventrally (0), or convex dorsoventrally (1).

• 293. Contour of articular surface for metacarpal V on metacarpal IV: concave dorsoventrally (0), or convex dorsoventrally (1).

• *294. Shape of articular surface for metacarpal V on metacarpal IV: ovate, proximodistal depth ≥ dorsoventral depth (0), ovate, compressed proximodistally (1), or triangular, tapers ventrally (2).

• 295. Lateral tubercle and proximal articulation on metacarpal V: lateral tubercle of metacarpal V at or below the level of proximal articular surface on metacarpal IV, articulation on metacarpal V for hamate fairly level (0), or metacarpal V forms peg-and-socket articulation with hamate, lateral tubercle of metacarpal V lies proximal to articular surface on metacarpal IV (1).

• 296. Cross-sectional shape of shaft of proximal and intermediate manual phalanges: width ≥ depth, shaft cylindrical or even slightly compressed dorsoventrally (0), or compressed mediolaterally, width < depth (1).

• 297. Lateral fossae for insertion of tendons on distal end of proximal manual phalanges: present (0), or absent (1).

• *298. Relative lengths of manual proximal (PP) and intermediate (IP) phalanges: nearly equivalent, ratio of IP/PP length <1.2 on all digits (0), proximal phalanx shorter on digits II and III (1), proximal phalanges shortened on digits III-V (2), or all proximal phalanges shortened (3).

• **299. Relative widths of proximal phalanges on digits II and IV: II > IV (0), II = IV (1), or II < IV (2).

• 300. Distal articulation of proximal phalanx on digit III: with paired distal condyles separate (0), or with paired distal condyles joined at their dorsoventral midpoint by a transverse eminence (1).

• *301. Proximal articulations of manual intermediate phalanges: visible in dorsal view, dorsal midline process does not extend as far proximally as basal tubera (0), not visible dorsally, dorsal midline process elongated proximally (1), or not visible dorsally, but dorsal midline process relatively short (2).

• **302. Size of manual ungual phalanx (UP) on digit I: roughly equal to UP V (0), reduced, smaller than UP V (1), or greatly reduced, <1/2 the length of UP V (2).

• 303. Size of ungual phalanx on digit III: not enlarged at all or only slightly enlarged relative to next largest ungual (0), or greatly enlarged, length >1.3 times that of next longest ungual (1).

• **304. Length of metatarsal III + proximal phalanx III + intermediate phalanx III + ungual phalanx III vs. tibial length: toes short, <70% tibial length (0), toes intermediate length, 70–85% of tibial length (1), or toes elongate, >90% tibial length (2).

• **305. Length of metacarpal III + proximal phalanx III + intermediate phalanx III + ungual phalanx III vs. humeral length: toes short, <60% humeral length (0), toes intermediate length, 70–80% of humeral length (1), or toes elongate, >80% humeral length (2).

• *306. Anterior border of nasal: with shallow notch (0), with deep notch forming elongated medial and lateral processes, with deep fossa on lateral process (1), or deeply concave, with elongated lateral process but medial process rudimentary or absent (2).

• 307. Nasal length: less than or equal to (0), or greater than one-third skull length (1).

• 308. Maxillonasal suture: subparallel (0), or convergent anterior to maxillofrontal junction (1).

• 309. Dorsal (facial) process of premaxilla: C-shaped, broad anteroposteriorly with distinct anteroventral and anterodorsal processes (0), or inclined posterodorsally (1).

• 310. Incisive foramen: within premaxilla (0), or between premaxilla and maxilla (1).

• 311. Size of palatal process of premaxilla: small or absent, if present, extending posteriorly between maxillae for a distance less than or equal to the maximum diameter of the incisive foramen (0), or elongate, extends posteriorly for a distance clearly greater than the maximum diameter of the incisive foramen (1).

• 312. Vomerine exposure on palate: absent (0), or present (1).

• 313. Maxilla with deep median longitudinal palatal concavity: absent (0), or present (1).

• 314. Alveolar sulcus on maxilla: absent (0), or present (1).

• 315. Maxilla with narrow posterior palatal process extending lateral to palatine: absent (0), or present (1).

• *316. Palatine foramina: multiple within palatine and maxilla (0), double within palatine (or posterior within palatine, anterior between palatine and maxilla) (1), single within maxilla (2), double within maxilla (3), or single large foramen within palatine, at times with smaller accessory foramina in palatine (4).

• *317. Anterolateral shelf of palatine: absent (0), lies posterior to (1), or medial to zygomatic process of maxilla (2).

• 318. Palatines within choanae: subparallel, widely separated by presphenoid and/or vomer (0), or converging anteriorly, nearly contacting (1).

• **319. Lacrimal bone and lacrimal foramen: both present (0), bone present, foramen absent (1), or both absent (2).

• 320. Lacrimal fenestra: absent (0), or present (1).

• 321. Ethmoidal foramen: within frontal (0), or between frontal and orbitosphenoid (1).

• **322. Orbitosphenoid/squamosal contact: absent (0), narrow, alisphenoid approximates frontal (1), or broad (2).

• 323. Small foramen posterior to optic foramen in orbitosphenoid: absent (0), or present (1).

• 324. Foramen rotundum and sphenorbital fissure: separate (0), or confluent, opening into same fossa (1).

• 325. Foramen subovale: absent (0), or present (1).

• 326. Foramen ovale: within alisphenoid (0), or between alisphenoid and squamosal (1).

• 327. Flange on pterygoid process behind foramen ovale: absent (0), or present (1).

• *328. Pterygoid hamulus: distal tip anterior to auditory tube (0), distal tip extends to level of fenestra vestibuli (1), distal tip extends to level of anterior rim of jugular foramen (2), or absent (3).

• 329. Pterygoid/ectotympanic contact: widely separated (0), or closely approximated or in contact (1).

• 330. Ectotympanic inflation: present (0), or absent (1).

• 331. Entotympanic or facet for entotympanic on basioccipital: present (0), or absent (1).

• **332. Course of internal carotid artery: transpromontorial (0), on the ventromedial edge of petrosal, lateral to entotympanic if present (1), or within or medial to entotympanic (2).

• 333. Auditory tube orientation: anteromedial (0), or medial (1).

• 334. Promontorium of petrosal: prominent, globose (0), or weakly developed, flat (1).

• **335. Course of facial nerve: open sulcus, crista parotica weak (0), open sulcus, crista parotica prominent (1), or closed canal (2).

• 336. Fusion between distal tip of tympanohyal and lateral surface of promontorium: absent (0), or present (1).

• 337. Position of fenestra cochleae: well separated from fenestra vestibuli, facing posteriorly and slightly laterally (0), or immediately next to fenestra vestibuli, facing laterally and slightly posteriorly (1).

• 338. Shielding of fenestra cochleae: open posteroventrally (0), or shielded posteroventrally by flange on petrosal (1).

• *339. Position of fossa incudis within epitympanic recess: in lateral wall, facing medially (0), in posterior wall, facing anteriorly (1), or in medial wall, facing laterally (2).

• 340. Squamosal participation in roof of epitympanic recess: absent or forming small part of lateral wall (0), or extensive, forming much of roof (1).

• 341. Surface contour of incudal facet of malleus: concave (0), or convex (1).

• **342. Orientation of mallear head/incudal facet: mallear head unrotated, incudal facet facing caudally and laterally (0), mallear head unrotated, incudal facet facing caudally and medially (1), or head rotated dorsad 90°, incudal facet facing dorsally, caudally and medially (2).

• *343. Incus: body gracile, may or may not be rectangular, crura elongated (0), body stout and rectangular, crura short (1), or body gracile with extreme elongation of crus longum, reduction of crus breve (2).

• 344. Stapedial columella: elongated, height nearly equal to or exceeding greatest width of footplate (0), or short, height much less than greatest width of footplate (1).

• 345. Mastoid/exoccipital contact: broad, squamosal well separated from lateral rim of jugular foramen (0), or narrow, squamosal approximates lateral rim of jugular foramen (1).

• 346. Vomer: underlies presphenoid in posterior nasopharynx, visible through choanae (0), or restricted to anterior nasal cavity, not visible from choanae (1).

• 347. Tympanic process of pterygoid: absent (0), or present (1). [Note: This feature was termed the “lateral wing of basisphenoid” in Gaudin and Wible (1999). In adult pangolins, the basisphenoid, alisphenoid, and pterygoid are fused. However, a juvenile Phataginus tricuspis (CM 41123) clearly shows this feature to be part of the pterygoid and not the basisphenoid.]

• **348. Tympanic process of basioccipital: absent or weak, does not reach level of promontorium (0), moderate, even with promontorium (1), or prominent, ventral to promontorium (2). [Note: This feature is termed the “basioccipital wing” in Gaudin and Wible (1999).]

• 349. Exoccipital constriction anterior to occipital condyle: present (0), or absent (1).

• 350. Foramen magnum shape: circular (0), or oval transversely (1).

• 351. Supraoccipital/parietals suture shape: subtends an angle greater than 90° (0), or less than or equal to 90° (1).

• 352. Nuchal crest: strongly developed (0), or rudimentary to absent (1).

• **353. Squamosal inflation posterior to external auditory meatus/porus acousticus: absent (0), weak, not clearly demarcated on external surface (1), or strong, with distinct lateral bulge on external surface (2).

• 354. Position of alisphenoid/squamosal suture: widely separated from (0), or approximating base of zygomatic process (1).

• *355. Orientation (and size) of zygomatic process of squamosal: laterally directed (0), ventrally directed, elongated (1), or ventrally directed, short (2).

• *356. Glenoid fossa: concave anteroposteriorly (0), convex anteroposteriorly (1), or flat anteroposteriorly (2).

• *357. Postglenoid foramen: on posterior aspect of zygomatic process (0), on lateral aspect of zygomatic process (1), posterior to zygomatic process (2), or absent (3).

• 358. Temporal lines: present (0), or absent (1).

• 359. Postorbital process: present (0), or absent (1).

• 360. Postorbital constriction: present (0), or rudimentary to absent (1).

• **361. Tentorial ossification: absent (0), present but weak, developed only inferiorly on petrosal (1), or present, strongly developed, extends to roof of cranial cavity (2).

• 362. Endocranial venous grooves: present (0), or absent (1).

• 363. Floor of middle cranial fossa: formed by sphenoid (0), or formed by squamosal (1).

• 364. Mandibular symphysis length: long, extends well posterior to level of anterior mental foramen (0), or short, ends at or near level of anterior mental foramen (1).

• **365. Mandibular symphysis shape in lateral view: with anterior convexity (0), straight profile (1), or with anterior concavity (2).

• **366. Anterodorsolaterally directed prongs on outer surface of mandibular symphyseal region: absent (0), incipiently developed as dorsolaterally extended folds of dorsal symphyseal margin (1), or well-developed, tooth-like conical prongs (2).

• 367. Elongated medial perforation in mandibular canal: absent (0), or present (1).

• 368. Elongated lateral perforation in mandibular canal: absent (0), or present (1).

• *369. Mandibular condyle: mediolaterally elongate (0), circular (1), or anteroposteriorly elongated (2).

• 370. Position of mandibular condyle: dorsal to (0), or at the level of the mandibular symphysis (1).

• **371. Coronoid process of mandible: present, well developed (0), present but strongly reduced in size (1), or absent (2).

• 372. Angular process of mandible: present (0), or absent (1).

• 373. Length of nasal/premaxillary vs. nasal maxillary suture: maxillonasal suture longer (0), or premaxillonasal suture longer (1).

• **374. Position of infraorbital foramen in lateral view: foramen situated toward back of maxilla, distance from maxillopalatine suture to infraorbital foramen <20% of the distance from maxillopalatine suture to maxillary/premaxillary suture (0), foramen displaced anteriorly, distance from maxillopalatine suture to infraorbital foramen ≥20%, <30% of the distance from maxillopalatine suture to maxillary/premaxillary suture (1), or foramen near midpoint of maxilla, distance from maxillopalatine suture to infraorbital foramen ≥30% of the distance from maxillopalatine suture to maxillary/premaxillary suture (2).

• **375. Position of maxillopalatine suture in ventral midline: posterior to maxillary foramen (0), at or slightly anterior to maxillary foramen (1), or well anterior to maxillary foramen (2).

• 376. Extension of orbital wing of palatine posterior to sphenorbital fissure: absent (0), or present (1).

• 377. Optic foramen size: small, diameter <2% greatest skull length (0), or large, diameter >3% (1).

• **378. Position of optic foramen relative to frontal/squamosal suture [or if absent lateral portion of frontal/parietal suture]: posterior to (0), approximately level with (1), or anterior to suture (2).

• 379. Position of foramen ovale: anterior to anterior edge of ectotympanic (0), or at level of anterior edge of ectotympanic (1).

• 380. Jugal: present (0), or absent (1).

• 381. Zygomatic arch: complete (0), or incomplete (1).

• 382. Frontal/parietal suture position: well anterior to glenoid, at or near postorbital constriction (0), at or posterior to glenoid, well posterior to postorbital constriction (1).

• 383. Alisphenoid/parietal contact: present (0), or absent (1).

• **384. Temporal fossa on braincase: large (0), slightly reduced in size (1), or strongly reduced (2).

• 385. Length of parietal/squamosal suture: <25% greatest skull length (0), or ≥25% (1).

• 386. Shape of zygomatic process of squamosal: elongated, uninflated, with blunt tip or elongated anterior sutural contact with jugal (0), or short, inflated at base, and pointed at tip (1).

• *387. Lateral exposure of mastoid and posttympanic process of squamosal: present (0), absent (1), or mastoid lateral exposure present, posttympanic process of squamosal absent (2).

• 388. Foramen for inferior petrosal sinus separate from jugular foramen: absent (0), or present (1).

• 389. Position of superior petrosal sinus: in groove between petrosal and tentorium (0), or perforates ventral portion of tentorium (1).

• *390. Position of jugular foramen and hypoglossal foramen: jugular foramen lateral to hypoglossal foramen, foramina well separated (0), jugular foramen lateral to hypoglossal foramen, foramina close, may even share a common fossa (1), jugular foramen anterior to or more medial than hypoglossal foramen, foramina well separated (2), or jugular and hypoglossal foramina confluent (3).

• 391. Epitympanic sinus between squamosal and petrosal: absent (0), or present (1).

• 392. Basicranial/basifacial axis: linear (0), or reflexed (1).

• *393. Teeth: normal mammalian dentition present (0), teeth present but reduced with large triangular canine but no incisors and only a few peg-like postcanine teeth (1), or teeth absent (2).

• 394. Depth of horizontal ramus of mandible: deep, >10% maximum mandibular length (0), or shallow, ≤10% (1).

• 395. Medial buttress on posterior portion of mandibular ramus: absent (0), or present (1).

Appendix 3

Data matrix.

The symbol ‘?’ represents either missing data or a case where a character is not applicable to a given taxon. The following symbols are used to represent character states in polymorphic taxa: a = (0, 1); b = (1, 2); c = (0, 1, 2); d = (1, 3); e = (0, 2); f = (3, 4); g = (2, 3); h = (0, 3); i = (1, 2, 3); j = (1, 4); k = (4, 5); m = (1, 5); n = (0, 1, 3). This data matrix has been deposited in MorphoBank and can be obtained at http://www.morphobank.org.

Table 3

Appendix 4

Distribution of apomorphies on the single MPT illustrated in Fig. 1. Characters shown in bold type are unambiguous synapomorphies; those in regular type are ambiguous synapomorphies. Those characters marked with a “^U” are unique apomorphies, as defined in the Results section.

Erinaceus <=> Nandinia + Pholidotamorpha:

4(3) <=> 4(0), 5(0) <=> 5(1), 7(0) <=> 7(1) ^U , 21(1) <=> 21(0), 25(1) <=> 25(2), 63(2) <=> 63(0), 74(2) <=> 74(1), 75(0) <=> 75(1), 76(1) <=> 76(0), 77(1) <=> 77(0), 93(1) <=> 93(2), 102(0) <=> 102(1), 107(1) <=> 107(0), 109(0) <=> 109(1), 110(2) <=> 110(0), 124(2) <=> 124(1), 135(0) <=> 135(1), 138(0) <=> 138(1), 142(2) <=> 142(0), 157(3) <=> 157(2), 158(2) <=> 158(1), 159(2) <=> 159(1), 161(2) <=> 161(3), 162(2) <=> 162(0), 165(2) <=> 165(1), 169(4) <=> 169(2), 175(1) <=> 175(0), 181(1) <=> 181(0), 186(0) <=> 186(1), 189(2) <=> 189(1), 201(3) <=> 201(2), 203(1) <=> 203(0), 207(3) <=> 207(2), 209(1) <=> 209(0), 221(0) <=> 221(1), 233(2) <=> 233(0), 237(0) <=> 237(1), 238(3) <=> 238(2), 239(2) <=> 239(1), 246(5) <=> 246(1), 251(1) <=> 251(0), 258(1) <=> 258(0), 264(2) <=> 264(0), 265(3) <=> 265(0), 272(0) <=> 272(1), 279(1) <=> 279(0), 293(2) <=> 293(0), 304(0) <=> 304(1), 305(0) <=> 305(1), 329(1) <=> 329(0), 332(0) <=> 332(1), 335(0) <=> 335(1), 342(0) <=> 342(1), 343(2) <=> 343(0), 350(2) <=> 350(1), 351(1) <=> 351(0), 354(0) <=> 354(1), 361(0) <=> 361(2), 376(1) <=> 376(0), 378(2) <=> 378(1), 385(1) <=> 385(0), 387(2) <=> 387(0).

Node 1. Pholidotamorpha:

10(1), 15(1), 19(1), 20(1), 45(1), 54(1), 79(0), 83(1), 86(0), 98(1), 106(0), 113(1), 145(0), 157(1), 165(0), 170(1), 176(1), 180(1), 196(0), 201(1), 206(0), 207(1), 212(1), 216(0), 217(0), 227(1), 230(1), 231(1), 240(1), 241(0), 260(1), 262(1), 269 (1), 273(1), 274(2), 306(0), 307(1), 309(0), 340(1) ^U , 373(0), 386(1), 390(0), 391(1).

Node 2. Palaeanodonta:

2(1), 71(0), 84(3), 89(0), 105(1), 109(2), 115(2) ^U , 116(0), 130(0), 155(0)^U, 208(0), 211(1), 215(1) ^U , 234(0), 246(3) ^U , 247(1), 248(0), 273(2), 280(0), 281(0), 282(2), 288(1) ^U , 290(2) ^U , 331(0), 356(0), 375(2), 393(1)^U, 395(1).

Node 3. Pholidota:

4(1), 37(1), 51(0), 81(1) ^U , 95(0), 100(4), 123(1), 147(0), 152(1) ^U , 155(1), 156(0), 202(0).

Node 4:

2(1), 3(0), 18(0), 19(0), 34(0), 45(2), 49(1), 56(0), 70(1), 103(0), 150(1)*^U, 160(1), 226(1), 236(0), 239(0), 240(2), 257(0), 305(2), 308(1), 314(1), 358(1)*^U, 366(1), 384(1), 392(1)*^U , 393(2)*^U , 394(1)*^U .

Node 5. Eupholidota:

11(1), 40(1), 63(1), 96(1), 129(1) ^U , 151(0), 155(2) ^U , 157(2), 168(2), 178(2), 192(1), 205(1)^U, 218(1), 235(1), 262(2), 263(1), 267(1), 286(0), 303(1), 313(1), 322(1), 370(1) ^U , 377(1), 384(2) ^U .

Convergences between Euromanis krebsi and Eomanis waldi: 3(1)^U.

Convergences between Euromanis krebsi and Node 5: 151(0).

Convergences between Euromanis krebsi and Eurotamandua: 68(0), 155(1)^U.

Convergences between Eurotamandua and Node 2, Palaeanodonta: 105(1) ^U , 247(1), 273(2), 282(2) ^U

Node 6. Manoidea:

1(1) ^U , 2(2), 4(0), 22(1), 33(2), 36(1)^U, 57(1), 65(1)^U, 67(2), 71(0), 80(1)^U, 95(1), 99(0), 106(1), 111(0), 117(1)^U, 128(1), 166(1) ^U , 167(1) ^U , 183(0), 186(0), 193(2), 197(1), 198(1) ^U , 199(1), 200(2), 208(0), 213(1), 215(2), 216(2), 225(0), 229(1), 234(0), 244(1), 249(1), 256(0), 274(1), 287(0), 306(1), 309(1), 320(1), 324(1)^U, 347(1)^U, 355(1), 366(2) ^U , 371(1), 374(0), 382(1), 383(1) ^U , 386(0).

Node 7. Patriomanidae:

2(0), 11(0), 14(0), 15(0), 42(1), 66(0), 74(0), 100(1) ^U , 109(0), 112(0) ^U , 118(1), 120(0), 130(0), 135(0), 146(0) ^U , 158(0), 163(0), 164(0), 180(0), 221(0), 228(0), 242(0), 261(0), 270(0), 288(0), 294(0), 297(0), 299(1), 305(1).

Synapomorphies between Necromanis and Patriomanidae (Node 7) on MPT #1:

3(0), 62(0), 66(0), 71(0), 74(0) ^U , 107(0), 109(0), 111(0), 151(0), 163(0), 164(0), 181(0), 183(0), 207(0) ^U , 208(0), 221(0), 225(0), 234(0), 235(1), 242(0), 251(0), 297(0).

Synapomorphies between Necromanis and Manidae (Node 8) on MPT #2:

2(2), 8(1), 11(1), 20(2), 22(1), 23(1) ^U , 42(2), 67(2), 92(1), 125(1), 128(2), 148(1) ^U , 200(2), 203(1), 215(2), 216(2), 229(1), 243(1), 248(2), 269(1), 303(1), 361(2), 372(1) ^U.

Node 8. Manidae:

3(2), 5(2) ^U , 6(1), 8(2) ^U , 9(1), 13(1), 21(1), 23(1), 24(1) ^U , 26(1), 27(1), 43(0), 49(2) ^U , 56(1), 59(1) ^U , 62(1), 63(2), 69(1), 70(2), 77(3), 89(2) ^U , 90(1), 91(1), 102(2) ^U , 103(2), 110(1), 125(2) ^U , 128(3), 131(1) ^U , 132(1) ^U , 136(1) ^U , 137(1), 140(1), 141(1), 142(1), 144(1) ^U , 145(1), 146(2), 148(1), 153(1) ^U , 154(1), 156(1), 157(3), 160(2), 171(1), 202(1), 203(1), 204(1) ^U , 209(1), 222(1), 223(1), 238(1), 244(2) ^U , 246(2) ^U , 248(2), 251(1), 252(1), 254(1) ^U , 257(1), 258(1), 260(2) ^U , 262(3), 264(1), 265(1), 266(1), 267(2), 275(1), 277(1), 282(1)^U, 284(1) ^U , 289(1) ^U , 290(1) ^U , 291(1), 293(1), 296(1) ^U , 298(3), 301(1) ^U , 302(1), 311(1), 316(2), 317(2), 319(1), 328(1), 333(1)^U , 334(1), 336(1), 337(1) ^U , 339(2) ^U , 341(1)^U , 346(1)^U , 352(1) ^U , 353(1), 356(1)^U , 360(1), 362(1) ^U , 363(1) ^U , 371(2), 372(1), 381(1), 387(1) ^U , 389(1) ^U .

Node 9. Maninae (genus Manis):

29(2), 30(2), 37(0), 41(2), 44(1), 46(3), 61(3), 70(3), 72(1) ^U , 74(2), 78(2), 79(2), 84(2), 88(1), 92(1), 93(3), 94(2), 107(1), 108(1) ^U , 118(1), 137(2), 149(2), 165(1), 169(3), 173(1), 182(2), 185(1) ^U , 188(1), 191(1), 194(1)^U , 214(0), 220(0), 224(1), 235(2), 241(1), 247(1), 255(1), 262(4) ^U , 279(1), 286(1), 294(2), 308(0), 319(2), 322(2) ^U , 335(2) ^U , 338(1), 343(1) ^U , 349(0), 364(0), 369(1), 377(0), 388(1), 390(2).

Node 10. M. crassicaudata + M. pentadactyla (Subgenus Manis):

12(2), 13(2), 16(1), 17(1), 19(1), 20(3), 58(1), 62(2), 67(1), 70(4), 71(1), 76(3), 104(2) ^U , 106(2), 110(2), 111(1), 114(0), 138(3), 139(3), 143(1), 151(1), 160(4), 161(0), 164(0), 168(3), 181(1), 211(1), 227(2), 230(0), 263(2), 271(0), 273(2), 276(2)^U , 277(2) ^U , 278(1), 283(1), 285(1) ^U , 291(2) ^U , 298(2), 300(1), 315(0) ^U , 350(0), 379(1).

Node 11. Smutsiinae (African manids):

20(2), 25(0), 28(1), 29(1), 30(1), 39(1), 47(1), 61(1), 76(2), 79(1), 82(1), 86(1), 90(2) ^U , 147(1), 161(1), 162(1), 177(0) ^U , 179(0), 181(1), 183(1), 184(1), 194(2)^U , 199(0), 201(2), 217(1), 225(1), 250(1), 261(2), 263(2), 268(1), 292(0), 295(1) ^U , 300(1), 312(1), 321(0), 330(0), 342(2) ^U , 344(1) ^U , 348(1), 369(2), 380(1).

Node 12. Phataginus:

4(2), 10(0), 11(2), 18(1), 19(1), 27(2), 31(2) ^U , 33(1) ^U , 38(1), 42(0), 45(1), 50(1) ^U , 52(1), 60(1) ^U , 62(2), 64(1) ^U , 68(0), 73(0), 74(2), 77(2), 83(2), 93(1), 98(0), 99(1), 113(2) ^U , 124(2), 126(1), 130(2) ^U , 149(0) ^U , 159(0), 165(2), 182(2), 187(0), 189(2), 190(0), 196(1), 207(2), 208(1), 210(1), 213(2), 216(3), 219(1) ^U , 227(0), 233(1), 234(1), 236(1), 239(1), 240(1), 241(2), 245(1), 256(1), 274(0), 275(2), 287(1), 302(2) ^U , 306(0), 316(3), 318(1), 323(1), 327(1), 329(1), 338(1), 350(0), 351(1), 353(2), 355(2), 367(1), 378(2), 385(1).

Node 13. Smutsia:

12(2), 18(2), 32(1), 34(1), 41(1), 44(1), 46(2), 48(0), 58(1), 71(1), 84(2), 87(1) ^U , 94(2), 107(1), 110(2), 111(1), 114(0), 119(1), 134(1), 137(2), 143(1), 145(3), 149(2), 151(1), 186(1), 193(1), 230(0), 231(2) ^U , 247(1), 269(0), 278(1), 283(1), 303(0), 310(0), 317(1), 319(2), 331(0), 335(0), 357(1), 368(1) ^U , 390 (1).