Details of datasets with inapplicable data

62 taxa, 242 characters, of which 78 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 4, 9, 30, 32, 33, 34, 35, 37, 38, 39, 40, 50, 52, 53, 55, 56, 63, 64, 65, 66, 67, 68, 69, 70, 72, 73, 74, 75, 76, 78, 79, 82, 83, 84, 85, 86, 87, 107, 108, 109, 111, 129, 144, 145, 148, 151, 152, 154, 157, 158, 159, 160, 161, 169, 170, 173, 175, 177, 178, 195, 196, 206, 216, 218, 223, 225, 227, 234

• Neomorphic: 36, 54, 59, 77, 81, 153, 156, 162, 226, 229

Details:

• N [153] absence (1) is secondary loss

76 taxa, 107 characters, of which 44 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 19, 20, 21, 26, 34, 40, 44, 51, 52, 55, 56, 64, 67, 82, 84, 91, 92, 97, 98, 99, 103

• Neomorphic: 13, 22, 25, 27, 28, 32, 33, 39, 45, 47, 48, 49, 50, 53, 54, 57, 58, 63, 70, 73, 75, 100, 101

Details:

• N [13] Second pair considered an additional state

• N [22] Articulation considered a novelty

• N [25] Triangularity considered derived

• N [27] Folding considered derived

• N [28] Fusion considered derived

• N [32] Additional pairs considered derived

• N [33] Articulation considered derived

• N [39] Inflation considered derived

• N [47] Articulation considered derived

• N [48] Articulation considered derived

• N [50] Differentiation considered derived

• N [53] Differentiation considered derived

• N [54] Alternation considered derived

• N [57] Articulation considered derived

• N [63] Restriction considered derived

• N [70] Differences considered derived

• N [73] Differentiation considered derived

• N [75] Differentiation considered derived

35 taxa, 50 characters, of which 39 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 2, 6, 7, 9, 13, 27, 37, 38, 39, 41, 44, 48, 49, 50

• Neomorphic: 15, 16, 17, 18, 19, 20, 21, 22, 23, 25, 26, 28, 29, 30, 31, 32, 33, 34, 35, 36, 40, 42, 43, 46, 47

Details:

• N [23] Character description missing; assumed N, following others of group III

23 taxa, 126 characters, of which 40 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 4, 7, 14, 33, 45, 48, 52, 54, 57, 66, 68, 69, 73, 76, 79, 80, 82, 88, 90, 103, 105, 109, 112, 117, 126

• Neomorphic: 16, 51, 70, 71, 72, 74, 75, 81, 83, 84, 85, 86, 87 ; with state 1 denoting absence: 78, 89

Details:

• X [1] post_stap_ramus / present reduced,

• X [2] sup_stap_ramus / present reduced,

• X [3] inf_stap_ramus / present reduced,

• T [4] opthart / IC SS IS EC,

• X [5] inf_alveolar / inf_stapedial ext_carotid,

• X [6] promontory_art / medial_cochlea lateral_cochlea,

• T [7] staped_bifurc / within_midear within_brcase,

• X [8] infraorb_in_cavepi / no yes,

• X [9] prox_ext_crtd / med_constrictor lat_constrictor,

• X [10] VNO_duct / into_NP_duct into_nasal_fossa,

• X [11] outer_bar / lat_closed lat_open,

• X [12] nasopal_duct / paired single,

• X [13] ‘NPduct cart-PRS’ / absent present,

• T [14] ‘NPduct cart-CNX’ / isolated cnx_parasept,

• X [15] ‘nasolac duct-PRS’ / present reduced,

• N [16] ‘nasolac duct-SHLD’ / unshielded shielded,

• X [17] papillary_cart / present reduced,

• X [18] VNO_vessels / large_superolat inconspic,

• X [19] Inf_nasal_sept / inf_parallel inf_rounded,

• X [20] zona_annularis / complete reduced,

• X [21] tymp_proc_basisphenoid / reduced present,

• X [22] piriform_fenestra / absent present,

• X [23] caudal_tymp_proc_petrosal / reduced shields_post_bulla,

• X [24] rostral_tymp_proc_petrosal / present nonapical,

• X [25] anterior_tympanic_roof / neither squamosal alisphenoid,

• X [26] epitympanic_recess / contin_w.midear enlarged,

• X [27] tubal_canal / not_closed_by_sph closed_by_sph,

• X [28] ‘ectotymp-PHAN’ / aphaneric semiphan phaneric,

• X [29] ‘ectotymp-EXPND’ / simple expanded,

• X [30] entotympanic / absent small contrib_bulla,

• X [31] infstap_for_alisph / absent present,

• X [32] ‘alisph canal-pres’ / absent present,

• T [33] ‘alisph canal-pos’ / lat_V contin_V,

• X [34] basisphenoid_pit / absent present,

• X [35] fenestrate_basioccip / no yes,

• X [36] suboptic_foramen / absent present,

• X [37] ‘mtuber-PM’ / reduced present hyper,

• X [38] ‘mtuber-SQ’ / reduced present,

• X [39] ‘SQtuber-jaw dist’ / short elongate,

• X [40] dual_ectopterygoid / no yes,

• X [41] maxilla_on_pteryg / no yes,

• X [42] postpal_spine / reduced present,

• X [43] petmast_position / expsd_posteriorly expsd_laterally not_expsd,

• X [44] ‘optic for-PRS’ / present absent,

• T [45] ‘optic for-SIZE’ / caliber_as_V smaller_V,

• X [46] ethmoid_for / anterior_SF within_SF,

• X [47] ‘sincan-PRS’ / present absent,

• T [48] ‘sincan-OPN’ / confl_w_SF ‘inbt SF-EF’ confl_w_EF,

• X [49] dorsal_palatine_for. / confl_w_sphpal separate,

• X [50] ‘entoglen-PRS’ / abs pres,

• N [51] ‘entoglen-JA’ / no_jaw_artic jaw_artic,

• T [52] ‘entoglen-COMP’ / all_squam half_alisph all_alisph,

• X [53] ‘lac formn-PRS’ / reduced present,

• T [54] ‘lac formn-OPN’ / open_post open_lat,

• X [55] orbital_wing_maxilla / no partial yes,

• X [56] ‘zygarch-PRS’ / complete incomplete,

• T [57] ‘zygarch-build’ / robust slender,

• X [58] origin_max_zygoma / M1_or_ant M2 M3,

• X [59] size_max_zygoma / posterolat_proj reduced,

• X [60] length_of_infraorb_canal / long short,

• X [61] Frontal_proportions / elongate even_w_parietal reduced,

• X [62] nasal_shape / broad narrow,

• X [63] nasal_length / ant_or_at_antorb post_to_antorb,

• X [64] lacrimal_flange / reduced prominent into_orbit,

• X [65] crista_galli / median_ridge flat,

• T [66] upper_ant_dent / ant_I_enlarged Is_equal,

• X [67] premaxillary_dentition / none 1ui 2ui 3ui 5ui,

• T [68] up.antmol_excl_Is / 5am 4am 3am ‘< 3am’,

• T [69] upper_Canine / premolariform bulbous trenchant,

• N [70] molariform_P3 / no yes,

• N [71] molariform_P4 / no yes,

• N [72] carnassial_P4 / no yes,

• T [73] metacone / present connate_w_paracone absent,

• N [74] protocone / present reduced, [Reduction is taken to represent the derived state]

• N [75] hypocone / present reduced, [Reduction is taken to represent the derived state]

• T [76] M1_parastyle / flush_wcing ant_proj,

• X [77] ‘M2/CBC’ / abs pres,

• I [78] ‘upper M3-PRS’ / pres abs,

• T [79] ‘upper M3-styshelf’ / short long,

• T [80] ‘upper M3-mstyle’ / lingual_meta buccal_meta,

• N [81] lower_ant_dent / ant_I_enlarged is_equal i2_enlarged, [Englargement is taken to represent the derived state]

• T [82] lower_antemolars / 8_antemolars 7_am 6_am 5_am 3_am,

• N [83] ‘molariform p/3’ / no_meta yes,

• N [84] ‘molariform p/4’ / no_meta yes,

• N [85] talonid_basin / present reduced,

• N [86] ‘m1-2 hypocnld’ / reduced present,

• N [87] ‘hypoflxd-pc occl’ / absent present,

• T [88] 2dary_hypoflxd_basin / closed open,

• I [89] ‘lower m3-PRS’ / present absent,

• T [90] ‘lower m3-LEN’ / m3_long ‘as m1-2’ m3_short,

• X [91] mental_foramnia / p4 p3 multiple m1,

• X [92] ‘mandib cond-LEN’ / narrow elongate,

• X [93] ‘mandib cond-NUM’ / single dual,

• X [94] condyle_height / at_toothrow above_toothrow,

• X [95] int_coronoid_fossa / _flat pocketed,

• X [96] ventral_crest_on_C2 / present absent,

• X [97] sacral_vertebrae / ‘2-4 verts’ ‘> 4 verts’,

• X [98] ‘verts in sacro-iliac art’ / s1_alone ‘s1-2’ ‘s1-3’,

• X [99] sacral_neural_spines / weak_or_not_fused fused abs_neural_spines,

• X [100] clavicles / present shortened absent,

• X [101] scapula / triangular elongate,

• X [102] ‘metacromion-PRS’ / present absent,

• T [103] ‘metacromion-shape’ / blunt elongate,

• X [104] ‘acromion-PRS’ / present absent,

• T [105] ‘acromion-shape’ / blunt elongate,

• X [106] ‘scapspine-len’ / short elongate,

• X [107] entepicondylar_for / present absent,

• X [108] medial_epicondyle / abs pres,

• T [109] ‘medepicond-shape’ / promin hypertr,

• X [110] olecranon_fenestra / absent thin_or_present,

• X [111] distal_ulna / present reduced,

• T [112] ulnocarpal_art / broad styloid_only,

• X [113] forearm_longbones / two three,

• X [114] iliac_blades / sag_or_rounded coronal,

• X [115] proximal_ilia / ‘cranial s-i joint’ ‘even s-i joint’,

• X [116] ‘iliopec tuber-PRS’ / weak prominent,

• T [117] ‘iliopec tuber-shape’ / extends_medially extends_inferiorly,

• X [118] internal_acetab_pelv_shape / smooth concave,

• X [119] pubic_symphysis / extensive reduced,

• X [120] pubis_length / as_isch_or_shorter longer_than_isch,

• X [121] angle_of_pubic_ramus / ‘> 15 angle’ parallel,

• X [122] 3rd_trochanter / proximal distal,

• X [123] ‘fusion of distal tibia-fib’ / no yes,

• X [124] distal_fibula / lateral_astrag socket_for_calc ball_for_astrag,

• X [125] calcaneal_tubercle / blunt elongate sep_fibular_facet,

• T [126] ‘interinc-gap’ / narrow wide,

67 taxa, 135 characters, of which 67 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 4, 6, 8, 9, 10, 16, 18, 19, 26, 34, 37, 40, 44, 46, 64, 69, 70, 74, 76, 79, 83, 84, 86, 88, 96, 99, 102, 104, 106, 121, 127, 131, 132, 135

• Neomorphic: 11, 12, 13, 14, 15, 17, 20, 21, 22, 23, 24, 25, 27, 28, 29, 30, 31, 32, 33, 35, 41, 43, 45, 47, 80, 81, 82, 85, 87, 100, 101, 103, 105

Details:

• X [1] Prostomium: (0) distinct; (1) completely fused to peristomium; (2) frontal edge fused to peristomium.

• X [2] Shape of prostomium: (0) lobed-ridge-shaped; (1) limited to palps; (2) conical.

• X [3] Prostomial palps: (0) absent; (1) present.

• T [4] Prostomial palps: (0) paired and simple; (1) multiple; (2) crown with radioles.

• X [5] Prostomial antennae: (0) absent; (1) present.

• T [6] Number and arrangement of prostomial antennae: (0) one in the middle; (1) two and lateral; (2) one median and two lateral.

• X [7] Prostomial eyes: (0) absent; (1) present.

• T [8] Radiolar lobes: (0) separated; (1) fused dorsally.

• T [9] Lobes: (0) semicircular or slightly involuted; (1) spiral.

• T [10] Number of radioles: (0) few (fewer than 3); (1) many (more than 3).

• N [11] Branching radioles: (0) absent; (1) present.

• N [12] Palmate membrane: (0) absent (or reduced); (1) present.

• N [13] Radiolar flanges: (0) absent; (1) present.

• N [14] Stylodes: (0) absent; (1) present.

• N [15] Radiolar skeleton axis cells: (0) absent; (1) present.

• T [16] Radiolar skeleton type: (0) one or two; (1) four; (2) more than four.

• N [17] Radiolar eyes: (0) absent; (1) present.

• T [18] Radiolar eyes arrangement: (0) ocelli; (1) unpaired terminal compound eyes on inner peduncle; (2) unpaired compound eyes on inner radiolar margin without peduncle; (3) paired compound eyes; (4) unpaired compound eyes on proximal outer margin of radioles; (5) cluster of eyes in the base of dorsalmost radioles.

• T [19] Pinnules: (0) alternating; (1) paired.

• N [20] Branchial hearts: (0) absent; (1) present.

• N [21] Dorsal basal flanges: (0) absent; (1) present.

• N [22] Joint in dorsal basal flanges: (0) absent; (1) present.

• N [23] Ventral basal flanges: (0) absent; (1) present.

• N [24] Ventral radiolar appendages: (0) absent; (1) present.

• N [25] Ventral radiolar filamentous appendages: (0) absent; (1) present.

• T [26] Ventral radiolar filamentous appendages type: (0) vascularized, unbranched; (1) vascularized, branched.

• N [27] Operculum: (0) absent; (1) present.

• N [28] Dorsal lips: (0) absent; (1) present.

• N [29] Dorsal radiolar appendages: (0) absent; (1) present.

• N [30] Radiolar appendage skeleton: (0) absent; (1) present.

• N [31] Dorsal pinnular appendages: (0) absent; (1) present.

• N [32] Ventral lips: (0) absent; (1) present.

• N [33] Ventral sacs: (0) absent; (1) present.

• T [34] Position of ventral sacs: (0) inside the branchial crown; (1) outside the branchial crown.

• N [35] Parallel lamellae: (0) absent; (1) present.

• X [36] Nuchal organs: (0) absent; (1) present.

• T [37] Type of nuchal organs: (0) pits or grooves; (1) two posterior projections; (2) caruncle.

• X [38] Peristomium: (0) reduced lips; (1) extended upper and lower lips; (2) one ring; (3) two rings.

• X [39] Peristomial palps: (0) absent; (1) present.

• T [40] Number of peristomial palps: (0) paired; (1) single; (2) multiple.

• N [41] Peristomial palp pinnules: (0) absent; (1) present.

• X [42] Peristomial eyes: (0) absent; (1) present.

• N [43] Anterior peristomial ring collar: (0) absent; (1) present.

• T [44] Anterior peristomial ring: (0) low, even in height; (1) reduced to a wide ventral triangular lobe; (2) reduced to a narrow ventral triangular lobe.

• N [45] Posterior peristomial ring collar: (0) absent; (1) present.

• T [46] Dorsal margin or posterior peristomial ring collar: (0) widely separated; (1) narrowly separated and fused to the faecal groove.

• N [47] Ventral incision of collar: (0) absent; (1) present.

• X [48] Tentacular cirri: (0) absent; (1) present.

• X [49] Obturaculum: (0) absent; (1) present.

• X [50] Vestimentum: (0) absent; (1) present.

• X [51] Frenulum: (0) absent; (1) present as a continuous ridge; (2) scattered plaques only.

• X [52] Trophosome: (0) absent; (1) present.

• X [53] Epidermal papillae: (0) absent; (1) present.

• X [54] Body covering or tunic: (0) absent; (1) present.

• X [55] Anterior end: (0) non-retractile; (1) retractile.

• X [56] Cephalic cage: (0) absent; (1) present.

• X [57] Glandular girdle on segment 2: (0) absent; (1) present.

• X [58] Thoracic membrane on anterior segments: (0) absent; (1) present.

• X [59] Ventral glandular areas: (0) absent or undifferentiated; (1) present.

• X [60] Interramal eyespots: (0) absent; (1) present.

• X [61] Chaetal inversion: (0) absent; (1) present.

• X [62] Inverted faecal groove: (0) absent; (1) present.

• X [63] Parapodia: (0) absent; (1) present.

• T [64] Parapodia: (0) similar rami; (1) noto- and neuropodia differing.

• X [65] Dorsal cirri: (0) absent; (1) present.

• X [66] Ventral cirri: (0) absent; (1) present.

• X [67] Aciculae: (0) absent; (1) present.

• X [68] Compound chaetae: (0) absent; (1) present.

• T [69] Thoracic chaetal fascicles (notopodia): (0) transverse; (1) longitudinal; (2) irregular.

• T [70] Thoracic neuropodia: (0) short, truncate cylinders; (1) lobe like; (2) transverse ridges (tori); (3) inconspicuous, chaetae from body wall; (4) low ridges; (5) conical.

• X [71] Notochaetae in first chaetiger: (0) absent; (1) present.

• X [72] Neurochaetae in first chaetiger: (0) absent; (1) present.

• X [73] Superior thoracic notochaetae: (0) absent; (1) present.

• T [74] Superior thoracic notochaetae shape: (0) elongate, narrowly hooded; (1) broadly hooded.

• X [75] Inferior thoracic chaetae: (0) absent; (1) present.

• T [76] Inferior thoracic notochaetae shape: (0) broadly hooded; (1) paleate; (2) spine-like; (3) narrowly hooded; (4) serrated capillaries; (5) hooks; (6) camerate chaetae; (7) capillary barred chaetae.

• X [77] Anterior row (or group) of inferior thoracic notochaetae: (0) absent; (1) bayonet.

• X [78] Thoracic uncini / hooks: (0) absent; (1) present.

• T [79] Arrangement of thoracic hooks/ uncini: (0) discrete single rows; (1) discrete double rows; (2) dense fields; (3) form rings.

• N [80] Rostrum / main fang: (0) absent; (1) present.

• N [81] Capitium: (0) absent; (1) present.

• N [82] Thoracic uncini teeth: (0) equal size; (1) unequal size. [Differentiated considered derived.]

• T [83] Rows of teeth on thoracic uncini: (0) 1–5; (1) more than 5 (numerous).

• T [84] Thoracic uncini with rows on only one tooth above main fang: (0) absent (various teeth per row); (1) present (one tooth per row).

• N [85] Thoracic uncini breast: (0) absent; (1) present.

• T [86] Development of thoracic uncini breast: (0) slightly developed; (1) well developed.

• N [87] Thoracic uncini handle / manubrium: (0) absent; (1) present.

• T [88] Length handles/ manubrium: (0) very long; (1) medium length; (2) short.

• X [89] Companion chaetae: (0) absent; (1) present (in all chaetigers); (2) present in some chaetigers.

• X [90] Number of abdominal chaetigers: (0) 3; (1) numerous.

• X [91] Opisthosome: (0) absent; (1) present.

• X [92] Abdominal neuropodia: (0) transverse ridges (tori); (1) conical lobes; (2) low, circular elevations; (3) straight from body wall; (4) several lobes.

• X [93] Abdominal neurochaetal fascicles: (0) transverse; (1) spiralled; (2) irregular.

• X [94] Superior row of abdominal neuropodial fascicles: (0) absent; (1) present.

• X [95] Inferior row of abdominal neurochaetal fascicles: (0) absent; (1) present.

• T [96] Superior row of abdominal neurochaetal fascicles: (0) narrowly hooded; (1) paleate; (2) spine-like; (3) broadly hooded; (4) capillaries; (5) stout and spinous.

• X [97] Inferior row of abdominal chaetae: (0) needle-like; (1) narrowly hooded; (2) spine-like; (3) broadly hooded; (4) denticulated capillaries; (5) flat-trumpet shaped; (6) stout and spinous; (7) paleate.

• X [98] Uncini in abdominal neuropodia: (0) absent; (1) present.

• T [99] Arrangement of abdominal hooks/ uncini: (0) single rows; (1) double rows; (2) dense fields.

• N [100] Abdominal uncini main fang: (0) absent; (1) present.

• N [101] Capitium: (0) absent; (1) present.

• T [102] Abdominal uncini number of rows: (0) few (1–5); (1) numerous (> 5).

• N [103] Uncini breast: (0) absent; (1) present.

• T [104] Abdominal uncini breast: (0) small, poorly developed; (1) well developed, expanded; (2) narrow, swelling; (3) proximal manubrium.

• N [105] Abdominal uncini handle: (0) absent; (1) present.

• T [106] Length of uncini handles: (0) short; (1) medium; (2) long.

• X [107] Anal depression: (0) absent; (1) present.

• X [108] Pygidial shape: (0) papilla; (1) rim; (2) bilobed.

• X [109] Pygidial cirri: (0) absent; (1) present.

• X [110] Pygidial eyes: (0) absent; (1) present.

• X [111] Caudal region: (0) absent; (1) present.

• X [112] Mouth: (0) axial proboscis; (1) simple tube non-eversible; (2) ventral buccal organ.

• X [113] Jaws: (0) absent; (1) present.

• X [114] Dorsolateral folds: (0) absent; (1) present.

• X [115] Gut: (0) as a tube with two openings; (1) occluded.

• X [116] Gut: (0) straight; (1) looped.

• X [117] Gular membrane: (0) absent; (1) present.

• X [118] Sexual mode: (0) gonochoric; (1) hermaphrodites.

• X [119] Gamete distribution: (0) abdomen; (1) thorax and abdomen; (2) thorax.

• X [120] Brooding: (0) absent; (1) present.

• T [121] Brooding: (0) intratubular; (1) extratubular.

• X [122] Spermiogenesis: (0) tetrads; (1) cytophore with more than 100 spermatids; (2) cytophore with fewer than 100 spermatids.

• X [123] Sperm packaging: (0) absent; (1) present.

• X [124] Larva: (0) trochophore; (1) mitraria.

• X [125] Larvae: (0) planktotrophic; (1) lecithotrophic.

• X [126] Lateral organs: (0) absent; (1) present.

• T [127] Segmental nephridia: (0) several; (1) single pair opening into first segment.

• X [128] Nephridia: (0) protonephridia; (1) metanephridia.

• X [129] Intraepidermal cord: (0) absent; (1) present.

• X [130] Branchiae: (0) absent; (1) present.

• T [131] Arrangement of branchiae: (0) dorsal, along body; (1) paired in anterior segments; (2) anterior cluster.

• T [132] Shape and number of of branchiae: (0) simple filaments; (1) multiple filaments; (2) laminate.

• X [133] Building organ: (0) absent; (1) present.

• X [134] Tube: (0) absent; (1) present.

• T [135] Tube material: (0) soft and mucous; (1) chitinous; (2) calcareous

64 taxa, 363 characters, of which 152 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 4, 7, 8, 9, 10, 13, 23, 27, 29, 30, 32, 33, 37, 38, 39, 43, 44, 46, 47, 49, 51, 52, 53, 62, 64, 67, 69, 70, 72, 78, 81, 84, 88, 90, 92, 93, 96, 97, 99, 104, 116, 125, 126, 129, 136, 138, 141, 145, 146, 149, 151, 155, 171, 176, 177, 178, 179, 183, 186, 189, 190, 191, 197, 198, 221, 223, 229, 234, 238, 242, 253, 261, 264, 267, 272, 274, 275, 276, 277, 278, 279, 280, 281, 282, 283, 285, 286, 288, 289, 291, 310, 326, 332, 333, 346

• Neomorphic: 17, 35, 42, 48, 50, 74, 75, 76, 82, 83, 89, 94, 100, 103, 110, 121, 137, 139, 156, 170, 173, 174, 185, 199, 202, 204, 210, 217, 233, 262, 263, 266, 269, 270, 273, 290, 292, 294, 299, 306, 308, 309, 311, 312, 314, 327, 328, 329, 330, 349, 356, 358, 359, 360, 363 ; with state 1 denoting absence: 87, 287

Details:

• T [8] Sculpture is taken as a transformation, rather than its addition in a region representing a novelty

• T [9] Sculpture is taken as a transformation, rather than its addition in a region representing a novelty

• T [10] Sculpture is taken as a transformation, rather than its addition in a region representing a novelty

• T [27] ‘contact’ is not a novel feature

• N [156] Loss indicates that presence (0) was original state

• N [290] Absence presumed to reflect a secondary loss

33 taxa, 50 characters, of which 19 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 3, 4, 11, 12, 13, 14, 16, 37, 38, 39, 40, 41, 42, 43, 44, 45, 46, 47, 48

• Neomorphic:

Details:

• X [1] Position of prostomium: (0) anterior to peristomium; (1) fused to peristomium.

• X [2] Shape of prostomium: (0) conical; (1) keel-shaped.

• T [3] Cephalic keel (prostomium): (0) slightly arched; (1) strongly arched; (2) flat.

• T [4] Anterior end of the prostomium: (0) narrow; (1) large, mushroom-shaped; (2) large, spade-shaped.

• X [5] Peristomium: (0) limited to lips; (1) forming the lips and post-prostomial rings.

• X [6] Nuchal grooves (paired slits): (0) dorso-lateral; (1) dorsal.

• X [7] Shape of nuchal grooves: (0) curved (elliptical); (1) slightly curved; (2) straight.

• X [8] Length of nuchal grooves in relation to keel: (0) shorter than keel; (1) of the same length as keel.

• X [9] Width of nuchal grooves in relation to keel: (0) narrow; (1) wide.

• X [10] Head: (0) cylindrical; (1) flattened, dorsally forming a cephalic plate.

• T [11] Lateral margin of cephalic plate: (0) thin ring; (1) expanded.

• T [12] Dorsal surface of cephalic plate: (0) smooth; (1) with transversal striae.

• T [13] Lateral lobules of cephalic plate: (0) entire; (1) auricular; (2) slightly notched.

• T [14] Latero-posterior surface of cephalic plate: (0) ornamented; (1) smooth.

• X [15] Neurochaetae of first chaetigers: (0) with rostrate uncinus on chaetigers 1–3; (1) with acicular spines on chaetigers 1–3.

• T [16] Number of hooks in chaetigers 1–3: (0) numerous; (1) up to 3 hooks per torus; (2) 3–7 hooks per torus.

• X [17] Notochaetae of anterior chaetigers: (0) capillaries; (1) with acicular spines.

• X [18] Formation of chaetae: (0) from the dorsal to the ventral side; (1) inverted.

• X [19] Number of row of uncini: (0) single; (1) double.

• X [20] Rostrate uncini: (0) with a hood; (1) no hood.

• X [21] Manubrioavicular uncini: (0) with long and curved shaft; (1) shaft very short.

• X [22] Capitium of neurochaetae: (0) with few or vestigial denticules; (1) numerous denticules.

• X [23] Subrostral process of rostrate uncini: (0) with barbules; (1) no barbules.

• X [24] Position of rows of notochaetae: (0) transversal along entire body; (1) longitudinal in posterior half of the body.

• X [25] Notopodial capillary chaetae: (0) smooth limbation; (1) with ornamented limbation.

• X [26] Other types of notopodial chaetae: (0) simple chaetae; (1) chaetae ruffled; (2) helicoidal chaetae.

• X [27] Number of chaetigerous segments: (0) 17–29; (1) from 30 to 70.

• X [28] Chaetiger 1: (0) smooth; (1) with a partial, globose collar; (2) complete, globose collar.

• X [29] Tori: (0) forming a narrow belt; (1) forming a globose belt behind the median segments.

• X [30] Length of median chaetigers: (0) isometric with anterior segments; (1) larger than the anterior segments.

• X [31] Posterior chaetigers: (0) not campanulate; (1) campanulate.

• X [32] Pre-anal segments: (0) numerous achaetigers; (1) small number of achaetigers.

• X [33] Lateral margins of the achaetigerous segments: (0) smooth; (1) with lobules.

• X [34] Length of the pre-anal achaetigers: (0) equal to lengths last chaetigerous segments; (1) decreasing in lengths.

• X [35] Shape of last pre-anal achaetigers: (0) similar to previous segments; (1) two thick rings.

• X [36] Pygidium: (0) conical; (1) flattened, forming a plate.

• T [37] Length of conical pygidium relative to adjacent segments: (0) isometric to last chaetigers; (1) larger than anterior segments.

• T [38] Surface of conical pygidium: (0) smooth; (1) with transversal striae.

• T [39] Color of conical pygidium: (0) unpigmented; (1) numerous dark rings.

• T [40] Disposition of rings: (0) closely spaced; (1) distantly spaced.

• T [41] Shape of posterior portion of conical pygidium: (0) straight; (1) top-shaped.

• T [42] Anal plate: (0) border formed by single ring; (1) border may be present only dorsally (petaloid-shaped); (2) well-developed dorsal ventral borders forming a anal cup.

• T [43] Lateral surface of anal plate: (0) entire; (1) with slits; (2) slits reduced to the borders of the anal cup.

• T [44] Posterior surface of dorsal border: (0) smooth (1) ornamented.

• T [45] Ornamentation of anal cup (ventral and dorsal borders): (0) cirri; (1) crenulations.

• T [46] Disposition of cirri of anal plate: (0) symmetrical; (1) asymmetrical.

• T [47] Size of ventral median cirrus: (0) similar to other cirri; (1) slightly longer than remaining cirri; (2) greatly elongate cirrus.

• T [48] Position of the anal pore within anal cup: (0) projected on a cone; (1) in level with the anal plate.

• X [49] Mechanism of closing anal pore: (0) exclusively muscular; (1) provided with a ventral valve.

• X [50] Position of anus: (0) terminal; (1) dorsal, to the side of the anal plate; (2) centre of anal plate

88 taxa, 220 characters, of which 18 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 7, 8, 26, 40, 41, 70, 124, 135, 137, 171, 176, 177, 200, 206, 219

• Neomorphic: 59, 107, 207

54 taxa, 131 characters, of which 41 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 8, 9, 20, 21, 23, 26, 29, 50, 51, 52, 53, 55, 58, 59, 61, 62, 63, 65, 66, 67, 68, 69, 70, 72, 73, 74, 77, 81, 93, 114, 115, 117

• Neomorphic: 24, 34, 42, 49, 54, 60, 64, 90 ; with state 1 denoting absence: 76

68 taxa, 186 characters, of which 87 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 1, 6, 8, 18, 21, 22, 24, 27, 33, 34, 37, 38, 83, 85, 86, 87, 90, 91, 94, 98, 99, 101, 103, 105, 106, 107, 109, 110, 111, 114, 115, 120, 122, 123, 126, 130, 151, 152, 153, 154, 156, 158, 160, 161, 162, 163, 164, 165, 169, 171, 172, 173, 174, 175, 179, 180

• Neomorphic: 19, 20, 25, 89, 92, 95, 96, 97, 116, 117, 118, 119, 121, 146, 155, 157, 159, 166, 167, 168, 170, 176, 177, 178 ; with state 1 denoting absence: 93, 100, 102, 104, 108, 112, 113

Details:

• T [1] Carotid_Notch_Bulla / absent present,

• X [2] Promontorial_Groove / present absent,

• X [3] Carotid_Foramen / absent present,

• X [4] Stapedial_Groove / present absent,

• X [5] Ramus_Sup._For. / present absent,

• T [6] POSITION_for_ram_sup / lateral anterolateral,

• X [7] size_pgf,

• T [8] Postglenoid_For. / squamosal suture absent,

• X [9] Mastoid_Foramen / present absent,

• X [10] Posttemporal_canal / present absent,

• X [11] Subarcuate_Fossa / present absent,

• X [12] Inflation_Tegmen_Tympani / uninflated inflated hyperinflated,

• X [13] Anterior_Proc_Petr / absent present,

• X [14] Tensor_Tympani_Fossa / shallow ‘circular pit, no groove’ ‘circular pit, anterior groove’ long_groove_between_tegmen_tymp,

• X [15] Stylomastoid_Foramen / complete incomplete,

• X [16] ‘Petrosal/Basioccpital’ / present absent,

• X [17] Ectotympanic_Bulla / ring ‘medial edge = bulla’,

• T [18] Bullar_Cavity / open cancellous,

• N [19] Pachyo_Bulla / absent present,

• N [20] Lat_furrow_Bulla / absent present,

• T [21] Inflation_Tymp._Bulla / absent same_level_as_condyles ‘present, ventral to condyles’,

• T [22] Bullar_Expansion / absent bulla_notched medial_expansion lateral_expansion stylohyal_surrounded,

• X [23] Stylohyal_Orient / ventral anteroventral,

• T [24] Art_Bulla_to_Squamosal / ‘contact broad, with medial pgp’ cirucular_facet_on_short_entogl contact_with_crest_of_entogleno co

• N [25] ‘Tymp/Exocc contact’ / absent present,

• X [26] Sigmoid_Process / absent present,

• T [27] Sigmoid_process_shape / thin_and_transverse_plate broad_and_flaring,

• X [28] EAM_Tympanic_Floor / absent present_but_short present_and_long,

• X [29] ‘Basioccipital crests (falcate p’ / absent present,

• X [30] Paroccipital_Proc. / short intermediate elongate,

• X [31] Post._Edge_Squamosal / flat sharply_upturned bears_dorsal_process,

• X [32] Lambdoidal_Crest / present absent,

• T [33] Sagittal_Crest_Height / absent_to_very_small ‘present, small’ ‘present, large’ ‘present, expanded’,

• T [34] Dorsal_Edge_Braincase / posterodorsal level posteroventral,

• X [35] Facial_Nerve_Sulcus / absent ant_wall_squamosal ant_wall_petrosal ant_wall_tympanic,

• X [36] Length_Mastoid_Proc / absent short long hypertrophied,

• T [37] Lateral_Exp._Mastoid / present absent,

• T [38] Mastoid_exp / exposed_externally not_exposed_posteriorly,

• X [39] Mastoid_angle,

• X [40] EAM_length,

• X [41] Postgl._Proc._Morph. / ridge rod,

• X [42] New_Glenoid / concave flat convex,

• X [43] Post._Edge_Glenoid_Fossa / flat notched_and_concave,

• X [44] Preglenoid_process / absent present,

• X [45] Posit_For_Ovale / ant_to_gf med_to_gf,

• X [46] Post._Edge_For._Ovale / alisphenoid petrosal squamosal,

• X [47] Zygoma_Height / narrow high dorsally_expanded,

• X [48] Zygoma_Direction / posterolateral posterior,

• X [49] Alisphenoid_Canal / absent present,

• X [50] Foramen_Rotundum / absent present,

• X [51] ‘Alisphenoid/Frontal’ / contact_present contact_absent,

• X [52] ‘Contact Front/Max’ / absent present,

• X [53] Supraorbital_Horns / absent ‘present, unbranched’ ‘present, branched’,

• X [54] Supraorbital_process / absent present,

• X [55] Postorbital_Pr._Jugal / absent ‘present, no frontal contact’ ‘present, contacts frontal’,

• X [56] Ventral_Edge_Orbit / projects_dorsally projects_laterally,

• X [57] Position_Orbit / Premolar first_M second_M third_M,

• X [58] Lacrimal_Tubercle / absent present,

• X [59] Lacrimal_foramina / two one highly_reduced absent,

• X [60] Facial_Lacrimal / restricted_to_edge facial_present expanded greatly_expanded,

• X [61] Lacrimal_Fossa / absent present,

• X [62] Facial_fenestra / absent present,

• X [63] Ant._Edge_Jugal / Premolar first_M second_M third_M,

• X [64] Elongation_Face / short long elongate,

• X [65] Infraorbital_Canal / over_M1_or_P4 between_P3P4 anterior_or_over_P3,

• X [66] lat_surface_maxilla,

• X [67] Posterior_Edge_Nasal / anterior_orbit over_orbit,

• X [68] Size_Palatine_Fissures / enlarged small absent,

• X [69] Palate_vaulting,

• X [70] Embrasure_pits / absent present,

• X [71] Post_margin_nares / anteriorover_canines between_P1_and_P2 posterior_to_P2,

• X [72] Angular_Process / curved dorsal_hook,

• X [73] Dentary_angle / level_with_ventral_edge below_ventral_edge,

• X [74] Mandibular_Foramen / small large,

• X [75] Coronoid_Process / short long elongate,

• X [76] Coronoid_Height / low high highest,

• X [77] Concavity_Mandible / absent present,

• X [78] Height_Dentary_Condyle / low high very_high,

• X [79] Dentary_Ramus / no_thickening ‘thickens from m1-m3’,

• X [80] Symphysis,

• X [81] ‘Upp Inc 1-2’ / present all_absent,

• X [82] Incisor_Alignment / transverse_arc ‘longitudinal, long prema’,

• T [83] Lower_Incisors / pointed spatulate ‘peg-shaped’ ‘tusk-like’,

• X [84] Upper_canine / short elongate,

• T [85] Lower_Canine / larger_than_incisors same_size smaller,

• T [86] Canine_shape / oval triangular,

• T [87] Canines,

• X [88] P1_presence / absent ‘present one-rooted’ ‘present two-rooted’,

• N [89] p1 / present absent,

• T [90] Lower_P1_Morph / low caniniform molariform,

• T [91] P3_roots / three two,

• N [92] p3_metaconid / absent present,

• I [93] P4_protocone / present absent,

• T [94] P4_paracone_height / equal_to_paracone_M1 greater_than_twice_M1_paracone,

• N [95] P4_metacone / absent present,

• N [96] P4_ectocingulum,

• N [97] p4_metaconid / absent present,

• T [98] dp4 / resembles_m1 6_cusped,

• T [99] M1_parastyle / absent weak strong,

• I [100] m1_metaconid / present absent,

• T [101] M2_metacone / subequal_to_paracone half_paracone highly_reduced,

• I [102] m2_metaconid / present absent,

• T [103] M3 / ‘present, larger than M2’ ‘present, approx. equal M2’ reduced absent,

• I [104] m3_metaconid / present absent,

• T [105] M3_Hypoconulid / long reduced absent,

• T [106] Para_posit,

• T [107] Stylar_shelves,

• I [108] Ectocingula / present absent,

• T [109] Mesostyle,

• T [110] Paraconule_Molars / present reduced absent,

• T [111] Cusp_number / 2_cusps 1_cusp no_cusps,

• I [112] protoloph / absent present,

• I [113] metaloph / absent present,

• T [114] lin_cing_molars / absent ‘continuous, mesial-distal’,

• T [115] ‘TEMP m1 parac/crist posit’ / cusp_or_crest_lingual ‘cusp or crest ant.betw pr-pa’,

• N [116] m1_m2_hypoconulid / absent present,

• N [117] Hypolophid / absent present,

• N [118] Metastylid / absent present,

• N [119] Entoconulid / absent present,

• T [120] protoconid / subequal_to_talonid twice_height_of_talonid,

• N [121] ant_low_lophs / absent present,

• T [122] Reen_groove / proximal absent distal,

• T [123] Talonid_basin / ‘basin, hy and ent present’ reduced absent,

• X [124] Occipital_Cond_Ridge / rounded clear_ridge,

• X [125] Occipital_Condyle_Edge / tapers_medially flared_laterally,

• T [126] Odontoid_Axis / anterior_peg ‘spout-like’ double_trough,

• X [127] Atlantoid_Facet_Axis / low high,

• X [128] Cerv_vert_length / short long very_long,

• X [129] Arterial_Canal_Cerv / post_open_external post_open_internal,

• T [130] ‘Art Sacral/Illium’ / broad narrow absent,

• X [131] Number_Sacrals / one two_or_three four five_or_six,

• X [132] Scapular_Spine / ‘large acromion, overhangs gleno’ small_acromion ‘large acromion, does not overha’,

• X [133] Supraspinatus,

• X [134] Entepicondyle / wide narrow,

• X [135] Entepi_foramen / present absent,

• X [136] Medial_Edge_Trochlea / restricted expanded,

• X [137] Intercondylar_Ridge / absent present,

• X [138] Olecranon_process / short long,

• X [139] Posterior_Edge_Ulna / convex straight concave,

• X [140] ‘Radius/Ulna’ / separate fused_distally fused_proximally,

• X [141] Proximal_radius / single_fossa two_fossae three_fossae,

• X [142] Distal_radius / single_fossa split_into_scaphoid_and_lunate,

• X [143] Centrale / present absent,

• X [144] ‘Magnum/Trapezoid’ / separate fused,

• X [145] Hand_Symm / between_2_and_3 mesaxonic paraxonic axis_along_4,

• N [146] ‘Metacarpal/Magnum’ / contact_present ‘absent, excl. 3 metacarpal’,

• X [147] Digit_2_Forelimb / long reduced,

• X [148] Width_2_Metacarpal / wide constricted very_narrow absent,

• X [149] Digit_5_Forelimb / long reduced,

• X [150] Width_5_Metacarpal / wide constricted very_narrow absent,

• T [151] Greater_troch / inferior_to_head_of_femur equal_to_head_of_femur above_head_of_femur,

• T [152] 3rd_troch / present highly_reduced absent,

• T [153] Distal_Femur / wide narrow,

• T [154] ‘Tibia/Fibula’ / separate fused_proximally fused_distally,

• N [155] Fibula_Completeness / complete incomplete, [Complete presumed ancestral]

• T [156] Proxi_astrag / flat grooved deeply_grooved,

• N [157] Astragalar_Canal / absent present,

• T [158] Navicular_Facet / convex flat ‘saddle-shaped’ trochleated,

• N [159] Cuboid_facet / absent ‘present-small’ present_transverse, [ Should be rewritten so treated as ordered ]

• T [160] Axes_of_Astragalus / obtuse parallel,

• T [161] Lateral_Astragalus / concave flat,

• T [162] Lateral_Proc_Astrag / present absent_indistinct,

• T [163] Sustentacular_facet / narrow wide,

• T [164] Sustentacular_Facet / separate_from_distal_facet partly_connected completely_connected,

• T [165] ‘Calcaneus/Cuboid’ / flat sharply_angled,

• N [166] ‘Nav/Cub fusion’ / unfused fused, [Fusion represents a derived state]

• N [167] ‘CuboNav/Ectocuneif’ / separate fused, [Fusion represents a derived state]

• N [168] ‘Ectocuneiform/Meso’ / separate fused, [Fusion represents a derived state]

• T [169] Pes / ‘mesaxonic-3’ paraxonic ‘mesaxonic-4’,

• N [170] Digit_1 / unreduced reduced highly_reduced,

• T [171] Digit_2_Hindlimb / long reduced_phalang reduced_metatars,

• T [172] Width_2_Metatarsal / wide constricted very_narrow medial_part_absent,

• T [173] Digit_5_Hindlimb / long reduced_phalanges,

• T [174] Width_5_Metatarsal / wide constricted very_narrow,

• T [175] Elong._Metatarsal_3 / short long elongate,

• N [176] Metapodial_Keels / absent present ‘present, extended anter’,

• N [177] ‘m3/m4 fusion’ / absent present,

• N [178] ‘3/4 Metatarsals’ / gully_absent present,

• T [179] Unguventral / curved_inferiorly straight,

• T [180] Ungdorsal / transversely_compressed ‘broad, symmetrical’ ‘broad, asymmetrical’,

• X [181] Hair / abundant absent_or_sparse,

• X [182] sebac_glands / present absent,

• X [183] fornix_expansion,

• X [184] omasum,

• X [185] Cavernous,

• X [186] Lung_bronchi,

78 taxa, 236 characters, of which 175 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 2, 6, 7, 10, 12, 13, 15, 16, 17, 19, 21, 22, 26, 28, 29, 30, 31, 35, 38, 39, 40, 41, 42, 43, 44, 45, 46, 47, 48, 49, 50, 54, 56, 57, 58, 59, 60, 61, 64, 65, 70, 71, 77, 78, 80, 82, 83, 84, 89, 90, 93, 96, 97, 99, 100, 106, 111, 114, 116, 138, 139, 144, 146, 150, 153, 154, 158, 161, 164, 182, 183, 184, 185, 192, 193, 197, 200, 202, 204, 216, 218, 219, 225, 228, 229, 231

• Neomorphic: 5, 8, 14, 18, 20, 23, 25, 27, 34, 36, 51, 53, 55, 63, 66, 67, 68, 69, 73, 74, 79, 81, 85, 86, 87, 88, 91, 92, 94, 98, 101, 102, 103, 104, 105, 107, 108, 112, 113, 120, 124, 125, 127, 129, 132, 133, 134, 141, 143, 145, 147, 149, 159, 160, 169, 172, 176, 180, 186, 187, 188, 189, 190, 191, 194, 195, 196, 198, 199, 203, 205, 206, 207, 209, 210, 211, 212, 220, 221, 222, 223, 224, 230, 233, 236 ; with state 1 denoting absence: 11, 24, 32, 181

Details:

• X [1] ‘[DFC12: 1] Tessellate prismatic calcified cartilage’: absent/ present,

• T [2] ‘Prismatic calcified cartilage’: ‘single layered’/ ‘multi-layered’,

• X [3] ‘[DFC12: 2] Perichondral bone’: present/ absent,

• X [4] ‘[DFC12: 3] Extensive endochondral ossification’: absent/ present,

• N [5] ‘Enamel(oid) present on dermal bones and scales’: absent/ present,

• T [6] Enamel: ‘single-layered’/ ‘multi-layered’,

• T [7] ‘Enamel layers’: ‘applied directly to one another (ganoine)’/ ‘separated by layers of dentine’,

• N [8] ‘Extensive pore canal network’: absent/ present,

• X [9] ‘[DFC10: 4] Dentinous tissue’: absent/ present,

• T [10] ‘[DFC10: 5] Dentine kind’: mesodentine/ semidentine/ orthodentine,

• I [11] ‘Bone cell lacunae in body scale bases’: present/ absent,

• T [12] ‘Main dentinous tissue forming fin spine’: osteodentine/ orthodentine,

• T [13] ‘[DFC12: 7] Longitudinal scale alignment in fin webs’: present/ absent,

• N [14] ‘Differentiated lepidotrichia’: absent/ present,

• T [15] ‘[DFC12: 8] Body scale growth pattern’: ‘comprising single odontode unit/generation (“monodontode”)’/ ‘comprising a complex of multipe odontode generations/units (“polyodontode”)’,

• T [16] ‘[DFC12: 9] Body scale growth concentric’: absent/ present,

• T [17] ‘Generations of odontodes’: buried/ ‘areally growing’/ resorbed,

• N [18] ‘[DFC12: 10] Body scales with peg-and-socket articulation’: absent/ present,

• T [19] ‘Scale peg’: broad/ narrow,

• N [20] ‘Anterodorsal process on scale’: absent/ ‘present’,

• T [21] ‘[DFC12: 11] Body scale profile’: ‘distinct crown and base demarcated by a constriction (“neck”)’/ flattened,

• T [22] ‘Profile of scales with constriction between crown and base’: ‘neck similar in width to crown’/ ‘neck greatly constricted, resulting in anvil-like shape’,

• N [23] ‘[DFC12: 12] Body scales with bulging base’: absent/ present,

• I [24] ‘[DFC12: 13] Body scales with flattened base’: present/ absent,

• N [25] ‘Basal pore in scales’: absent/ present,

• T [26] ‘[DFC12: 14] Flank scale alignment’: ‘vertical rows oblique rows or hexagonal’/ ‘rhombic packing’/ disorganised,

• N [27] ‘Scute-like ridge scales (basal fulcra)’: absent/ present,

• T [28] ‘[DFC12: 15] Sensory line canal’: ‘perforates scales’/ ‘passes between scales’/ ‘C-shaped scales’,

• T [29] ‘Dermal ornamentation’: smooth/ ‘parallel, vermiform ridges’/ ‘concentric ridges’/ tuberculate,

• T [30] ‘[DFC12: 16] Sensory line network’: ‘preserved as open grooves (sulci) in dermal bones’/ ‘sensory lines pass through canals in dermal bones (open as pores)’,

• T [31] ‘Sensory canals/grooves’: ‘contained within the thickness of dermal bones’/ ‘contained in prominent ridges on visceral surface of bone’,

• I [32] ‘[DFC 17] Jugal portion of infraorbital canal joins supramaxillary canal’: present/ absent,

• X [33] ‘[DFC 18] Dermal skull roof’: ‘includes large dermal plates’/ ‘consists of undifferentiated plates or tesserae’,

• N [34] ‘Anterior pit line of dermal skull roof’: absent/ present,

• T [35] ‘[DFC 19] Tessera morphology’: ‘large interlocking polygonal plates’/ ‘microsquamose, not larger than body squamation’,

• N [36] ‘Cranial spines’: absent/ ‘present, multicuspid’/ ‘present, monocuspid’,

• X [37] ‘[DFC 20] Extent of dermatocranial cover’: complete/ ‘incomplete (limited to skull roof)’,

• T[?] [38] ‘[DFC 21] Openings for endolymphatic ducts in dermal skull roof’: present/ absent,

• T [39] ‘[DFC 22] Endolymphatic ducts with oblique course through dermal skull bones’: absent/ present,

• T [40] ‘Endolymphatic duct relationship to median skull roof bone (i.e. nuchal plate)’: ‘within median bone’/ ‘on bones flanking the median bone (e.g. paranuchals)’,

• T[?] [41] ‘[DFC 25] Pineal opening perforation in dermal skull roof’: present/ absent,

• T [42] ‘Dermal plate associated with pineal eminence or foramen’: ‘contributes to orbital margin’/ ‘plate bordered laterally by skull roofing bones’,

• T [43] ‘[DFC 23] Series of paired median skull roofing bones that meet at the dorsal midline of the skull (rectilinear skull roof pattern)’: absent/ present,

• T? [44] ‘Broad supraorbital vaults’: absent/ present,

• T? [45] ‘Median commisure between supraorbital sensory lines’: absent/ present,

• T? [46] ‘Dermal cranial joint at level of sphenoid-otic junction’: absent/ present,

• T [47] ‘Otic canal extends through postparietals’: absent/ present,

• T [48] ‘Number of bones of skull roof lateral to postparietals’: two/ one,

• T [49] ‘Suture between paired skull roofing bones (centrals of placoderms; postparietals of osteichthyans)’: straight/ sinusoidal,

• T [50] ‘Medial processes of paranuchal wrapping posterolateral corners of nuchal plate’: absent/ present/ ‘paranuchals precluded from nuchal by centrals’/ ‘no median posterior skull roof bone’,

• N [51] ‘Paired pits on ventral surface of nuchal plate’: absent/ present,

• X [52] ‘Sclerotic ring’: absent/ present,

• N [53] ‘[DFC 24] Consolidated cheek plates’: absent/ present,

• T [54] ‘Cheek plate’: undivided/ ‘divided (i.e., squamosal and preopercular)’,

• N [55] ‘Subsquamosals in taxa with divided cheek’: absent/ present,

• T [56] ‘Preopercular shape’: rhombic/ ‘bar-shaped’,

• T [57] ‘Vertical canal associated with preopercular/suborbital canal’: absent/ present,

• T [58] ‘[DFC 26] Enlarged postorbital tessera separate from orbital series’: absent/ present,

• T [59] ‘Extent of maxilla along cheek’: ‘to posterior margin of cheek’/ ‘cheek bones exclude maxilla from posterior margin of cheek’,

• T [60] ‘Dermal neck joint’: overlap/ ginglymoid,

• T [61] ‘[DFC 15] Sensory line scales/plates on head’: unspecialized/ ‘apposed growth’/ ‘paralleling canal’/ ‘semicylindrical C-shaped ring scales’,

• X [62] ‘[DFC 27] Bony hyoidean gill-cover series (branchiostegals)’: absent/ present,

• N [63] ‘[DFC 28] Branchiostegal plate series along ventral margin of lower jaw’: absent/ present,

• T [64] ‘[DFC 29] Branchiostegal ossifications’: ‘plate-like’/ ‘narrow and ribbon-like’/ filamentous,

• T [65] ‘[DFC 30] Branchiostegal ossifications’: ornamented/ unornamented,

• N [66] ‘[DFC 31] Imbricated branchiostegal ossifications’: absent/ present,

• N [67] ‘Median gular’: absent/ present,

• N [68] ‘Lateral gular’: absent/ present,

• N [69] ‘[DFC 33] Opercular (submarginal) ossification’: absent/ present,

• T [70] ‘[DFC 34] Shape of opercular (submarginal) ossification’: ‘broad plate that tapers towards its proximal end’/ ‘narrow, rod-shaped’,

• T [71] ‘[DFC 36] Size of lateral gular plates’: ‘extending most of length of the lower jaw’/ ‘restricted to the anterior third of the jaw (no longer than the width of three or four branchiostegals)’,

• X [72] ‘Gill arches’: ‘largely restricted to region under braincase’/ ‘extend far posterior to braincase’,

• N? [73] ‘[DFC 37] Basihyal’: present/ absent,

• N? [74] ‘[DFC 38] Interhyal’: absent/ present,

• X [75] Hypohyal: absent/ present,

• X [76] ‘Endoskeletal urohyal’: absent/ present,

• T [77] ‘[DFC 39] Oral dermal tubercles borne on jaw cartilages or at margins of the mouth’: absent/ present,

• T [78] ‘[DFC 39] Oral dermal tubercles patterned in organised rows (teeth)’: absent/ present,

• N [79] ‘Enamel(oid) on teeth’: absent/ present,

• T [80] ‘Cap of enameloid restricted to upper part of teeth (acrodin)’: absent/ present,

• N [81] ‘[DFC 40] Tooth whorls’: absent/ present,

• T [82] ‘[DFC 41] Bases of tooth whorls’: ‘single, continuous plate’/ ‘some or all whorls consist of separate tooth units’,

• T [83] ‘[DFC42] Distribution of tooth whorls’: ‘entire length of tooth row’/ ‘restricted to symphysial region’,

• T [84] ‘Distribution of tooth whorls’: ‘upper and lower jaws’/ ‘lower jaws only’/ ‘upper jaws only’,

• N [85] ‘[DFC 43] Teeth ankylosed to dermal bones’: absent/ present,

• N [86] Plicidentine: absent/ present,

• N [87] ‘[DFC44] Dermal jaw plates on biting surface of jaw cartilages’: absent/ present,

• N [88] ‘[DFC 45] Maxillary and dentary marginal bones of mouth’: absent/ present,

• T [89] Premaxilla: ‘extends under orbit’/ ‘restricted anterior to orbit’,

• T [90] ‘Maxilla shape’: ‘splint-shaped’/ ‘cleaver-shaped’,

• N [91] ‘Pair of tooth plates (anterior supragnathals or vomers) on ethmoidal plate’: absent/ present,

• N [92] ‘Strong posterior flexion of dentary symphysis’: absent/ present,

• T [93] ‘Extent of infradentaries’: ‘along much of ventral margin of dentary’/ ‘restricted to posterior half of dentary’,

• N [94] ‘Coronoid fangs’: absent/ present,

• X [95] ‘Position of upper mandibular arch cartilage (and associated cheek plate where present)’: ‘entirely suborbital’/ ‘with a postorbital extension’,

• T [96] ‘Position of mandibular arch articulations’: terminal/ subterminal,

• T [97] ‘Autopalatine and quadrate’: comineralized/ ‘separate mineralizations’,

• N [98] ‘[DFC 46] Large otic process of the palatoquadrate’: absent/ present,

• T [99] ‘[DFC 47] Insertion area for jaw adductor muscles on palatoquadrate’: ‘ventral or medial’/ lateral,

• T [100] ‘Palatoquadrate relationship to dermal cheek bones’: ‘broad articulation’/ ‘articulation narrow and restricted’,

• N [101] ‘Palatoquadrate fused with neurocranium’: absent/ present,

• N [102] ‘[DFC 48] Oblique ridge or groove along medial face of palatoquadrate’: absent/ present,

• N [103] ‘[DFC 49] Fenestration of palatoquadrate at basipterygoid articulation’: absent/ present,

• N [104] ‘[DFC 50] Perforate or fenestrate anterodorsal (metapterygoid) portion of palatoquadrate’: absent/ present,

• N [105] ‘[DFC 51] Pronounced dorsal process on Meckelian bone or cartilage’: absent/ present,

• T [106] ‘Number of coronoids’: ‘four or more’/ ‘three or fewer’,

• N [107] ‘[DFC 52] Preglenoid process’: absent/ present,

• N [108] ‘[DFC 53] Jaw articulation located on rearmost extremity of mandible’: absent/ present,

• X [109] ‘[DFC 54] Precerebral fontanelle’: absent/ present,

• X [110] ‘[DFC 55] Median dermal bone of palate (parasphenoid)’: absent/ present,

• T [111] Parasphenoid: ‘lozenge-shaped’/ ‘splint-shaped’,

• N [112] ‘[Z13 241] Multifid anterior margin of parasphenoid denticle plate’: absent/ present,

• N [113] ‘Enlarged ascending processes of parasphenoid’: absent/ present,

• T [114] ‘Buccohypophysial canal in parasphenoid’: single/ paired,

• X [115] ‘[DFC 56] Nasal opening(s)’: ‘dorsal, placed between orbits’/ ‘ventral and anterior to orbit’,

• T [116] ‘Posterior nostril’: ‘separated from orbital fenestra’/ ‘confluent with orbital fenestra’,

• X [117] ‘[DFC 57] Olfactory tracts’: ‘short, with olfactory capsules situated close to telencephalon cavity’/ ‘elongate and tubular (much longer than wide)’,

• X [118] ‘[DFC 58] Prominent pre-orbital rostral expansion of the neurocranium’: present/ absent,

• X [119] ‘[DFC 59] Pronounced sub-ethmoidal keel’: absent/ present,

• N [120] ‘Internasal vacuities’: absent/ present,

• X [121] ‘Discrete division of the ethmoid and more posterior braincase at the level of the optic tract canal’: absent/ present,

• X [122] ‘[DFC 60] Position of myodome for superior oblique eye muscles’: ‘posterior and dorsal to foramen for nerve II’/ ‘anterior and dorsal to foramen’,

• X [123] ‘[DFC 61] Endoskeletal intracranial joint’: absent/ present,

• N [124] ‘[DFC 62] Spiracular groove on basicranial surface’: absent/ present,

• N [125] ‘Transverse otic process’: present/ absent,

• X [126] ‘Jugular canal’: ‘long (invested in otic region along length of skeletal labyrinth)’/ ‘short (restricted to region anterior of skeletal labyrinth)’/ ‘absent (jugular vein uninvested in otic region)’,

• N [127] ‘[DFC 63] Spiracular groove on lateral commissure’: absent/ present,

• X [128] ‘[DFC 64] Subpituitary fenestra’: absent/ present,

• N [129] ‘[DFC 65] Supraorbital shelf broad with convex lateral margin’: absent/ present,

• X [130] ‘[DFC 66] Orbit dorsal or facing dorsolaterally, surrounded laterally by endocranium’: present/ absent,

• X [131] ‘Eyestalk attachment area’: absent/ present,

• N [132] ‘Postorbital process’: absent/ present,

• N [133] ‘Canal for jugular in postorbital process’: absent/ present,

• N [134] ‘Series of perforations for innervation of supraorbital sensory canal in supraorbital shelf’: absent/ present,

• X [135] ‘[DFC 67] Extended prehypophysial portion of sphenoid’: absent/ present,

• X [136] ‘[DFC 68] Narrow interorbital septum’: absent/ present,

• X [137] ‘[DFC 69] The main trunk of facial nerve (N. VII)’: ‘elongate and passes anterolaterally through orbital floor’/ ‘stout, divides within otic capsule at the level of the transverse otic wall’,

• T [138] ‘[DFC 70] Course of hyoid ramus of facial nerve (N. VII) relative to jugular canal’: ‘traverses jugular canal, with separate exit in otic region’/ ‘intersects jugular canal, with exit through posterior jugular foramen’,

• T [139] ‘[DFC 71] Glossopharyngeal nerve (N. IX) exit’: ‘foramen situated posteroventral to otic capsule and anterior to metotic fissure’/ ‘through metotic fissure’,

• X [140] ‘[DFC 74] Relationship of cranial endocavity to basisphenoid’: ‘endocavity occupies full depth of sphenoid’/ ‘enodcavity dorsally restricted’,

• N [141] ‘Subcranial ridges’: absent/ present,

• X [142] ‘[DFC 75] Ascending basisphenoid pillar pierced by common internal carotid’: absent/ present,

• N [143] ‘[DFC 77] Canal for lateral dorsal aorta within basicranial cartilage’: absent/ present,

• T [144] ‘[DFC 78] Entrance of internal carotids’: ‘through separate openings flanking the hypophyseal opening or recess’/ ‘through a common opening at the central midline of the basicranium’,

• N [145] ‘[DFC 79] Canal for efferent pseudobranchial artery within basicranial cartilage’: absent/ present,

• T [146] ‘[DFC 80] Position of basal/basipterygoid articulation’: ‘same anteroposterior level as hypophysial opening’/ ‘anterior to hypophysial opening’,

• N [147] ‘[DFC: 81] Articulation between neurocanium and palatoquadrate posterodorsal to orbit (suprapterygoid articulation)’: absent/ present,

• X [148] ‘[DFC 82] Labyrinth cavity’: ‘separated from the main neurocranial cavity by a cartilaginous or ossified capsular wall’/ ‘skeletal capsular wall absent’,

• N [149] ‘[DFC 83] Basipterygoid process (basal articulation) with vertically oriented component’: absent/ present,

• T [150] ‘[DFC 84] Pituitary vein canal’: ‘dorsal to level of basipterygoid process’/ ‘flanked posteriorly by basipterygoid process’,

• X [151] ‘[DFC 85] External (horizontal) semicircular canal’: absent/ present,

• X [152] ‘[DFC 86] Sinus superior’: ‘absent or indistinguishable from union of anterior and posterior canals with saccular chamber’/ present,

• T [153] ‘[DFC 87] External (horizontal) semicircular canal’: ‘joins the vestibular region dorsal to posterior ampulla’/ ‘joins level with posterior ampulla’,

• T [154] ‘Horizontal semicircular canal in dorsal view’: ‘medial to path of jugular vein’/ ‘dorsal to jugular vein’,

• X [155] ‘Lateral cranial canal’: absent/ present,

• X [156] ‘[DFC 88] Trigemino-facial recess’: absent/ present,

• X [157] ‘[DFC 89] Posterior dorsal fontanelle’: absent/ present,

• T [158] ‘[DFC 90] Shape of posterior dorsal fontanelle’: ‘approximately as long as broad’/ ‘much longer than wide, slot-shaped’,

• N [159] ‘Synotic tectum’: absent/ present,

• N [160] ‘[DFC 91] Dorsal ridge’: absent/ present,

• T [161] ‘Shape of median dorsal ridge anterior to endolymphatic fossa’: ‘developed as a squared-off ridge or otherwise ungrooved’/ ‘bears a midline groove’,

• X [162] ‘[DFC 92] Endolymphatic ducts in neurocranium’: ‘posteriodorsally angled tubes’/ ‘tubes oriented vertically through median endolymphatic fossa’,

• X [163] ‘[DFC 95] Position of hyomandibula articulation on neurocranium’: ‘below or anterior to orbit, on ventrolateral angle of braincase’/ ‘on otic capsule, posterior to orbit’,

• T [164] ‘[DFC 93, in part] Position of hyomandibula articulation relative to structure of skeletal labyrinth’: ‘anterior or lateral to skeletal labyrinth’/ ‘at level of posterior semicircular canal’,

• X [165] ‘[Z13: 227] Hyoid arch articulation on braincase’: single/ double,

• X [166] ‘Branchial ridges’: present/ ‘reduced to vagal process’/ ‘absent (articulation made with bare cranial wall)’,

• X [167] ‘Craniospinal process’: absent/ present,

• X [168] ‘[DFC 96] Ventral cranial fissure’: absent/ present,

• N [169] ‘Basicranial fenestra’: absent/ present,

• X [170] ‘[DFC 97] Metotic (otic-occipital) fissure’: absent/ present,

• X [171] ‘[DFC 98] Vestibular fontanelle’: absent/ present,

• N [172] ‘[DFC 99] Occipital arch wedged in between otic capsules’: absent/ present,

• X [173] ‘[DFC 100] Spino-occipital nerve foramina’: ‘two or more, aligned horizontally’/ ‘one or two, dorsoventrally offset’,

• X [174] ‘[DFC 101] Ventral notch between parachordals’: ‘present or entirely unfused’/ absent,

• X [175] ‘[DFC 102] Parachordal shape’: ‘forming a broad, flat surface as wide as the otic capsules’/ ‘mediolaterally constricted relative to the otic capsules’,

• N [176] ‘Stalk-shaped parachordal/occipital region’: absent/ present,

• X [177] ‘Paired occipital facets’: absent/ present,

• X [178] ‘Size of aperture to notochordal canal’: ‘much smaller than foramen magnum’/ ‘as large, or larger, than foramen magnum’,

• X [179] ‘Canal for median dorsal aorta within basicranium’: absent/ present,

• N [180] ‘[DFC 103] Hypotic lamina (and dorsally directed glossopharyngeal canal)’: absent/ present,

• I [181] ‘[DFC 104] Macromeric dermal shoulder girdle’: present/ absent,

• T [182] ‘[DFC 105] Dermal shoulder girdle composition’: ‘ventral and dorsal (scapular) components’/ ‘ventral components only’,

• T [183] ‘[CA96: 115] Shape of dorsal blade of dermal shoulder girdle (either cleithrum or anterolateral plate)’: spatulate/ pointed,

• T [184] ‘[DFC 106] Dermal shoulder girdle forming a complete ring around the trunk’: present/ absent,

• T [185] ‘[DFC 107] Pectoral fenestra completely encircled by dermal shoulder armour’: present/ absent,

• N [186] ‘[DFC 108] Median dorsal plate’: absent/ present,

• N [187] ‘Posterior dorsolateral (PDL) plate or equivalent’: absent/ present,

• N [188] ‘[DFC 109] Pronounced internal crista (keel) on median dorsal surface of shoulder girdle’: absent/ present,

• N [189] ‘[DFC 109] Crista internalis of dermal shoulder girdle’: absent/ present,

• N [190] ‘Scapular infundibulum’: absent/ present,

• N [191] ‘[DFC 110] Scapular process of shoulder endoskeleton’: absent/ present,

• T [192] ‘[DFC 111] Ventral margin of separate scapular ossification’: horizontal/ ‘deeply angled’,

• T [193] ‘[DFC 112] Cross sectional shape of scapular process’: ‘flattened or strongly ovate’/ subcircular,

• N [194] ‘[DFC 113] Flange on trailing edge of scapulocoracoid’: absent/ present,

• N [195] ‘[DFC 114] Scapular process with posterodorsal angle’: absent/ present,

• N [196] ‘[DFC 115] Endoskeletal postbranchial lamina on scapular process’: present/ absent,

• T [197] ‘[DFC 116] Mineralisation of internal surface of scapular blade’: ‘mineralised all around’/ ‘unmineralised on internal face forming a hemicylindrical cross-section’,

• N [198] ‘[DFC 117] Coracoid process’: absent/ present,

• N [199] ‘[DFC 118] Procoracoid mineralisation’: absent/ present,

• T [200] ‘[DFC 119] Fin base articulation on scapulocoracoid’: ‘deeper than wide (stenobasal)’/ ‘wider than deep (eurybasal)’,

• X [201] ‘Pectoral fin articulation’: monobasal/ polybasal,

• T [202] ‘Number of basals in polybasal pectoral fins’: ‘three or more’/ two,

• N [203] ‘Branching radials in paired fins’: absent/ present,

• T [204] ‘Number of mesomeres in metapterygial axis’: ‘five or fewer’/ ‘seven or more’,

• N [205] ‘Biserial pectoral fin endoskeleton’: absent/ present,

• N [206] ‘[DFC 120] Perforate propterygium’: absent/ present,

• N [207] ‘Filamentous extension of pectoral fin from axillary region’: absent/ present,

• X [208] ‘[DFC 121] Pelvic fins’: absent/ present,

• N [209] ‘[DFC 122] Pelvic claspers’: absent/ present,

• N [210] ‘[DFC 123] Dermal pelvic clasper ossifications’: absent/ present,

• N [211] ‘[DFC 124] Pectoral fins covered in macromeric dermal armour’: absent/ present,

• N [212] ‘[DFC 125] Pectoral fin base has large, hemispherical dermal component’: absent/ present,

• X [213] ‘[DFC 126] Dorsal fin spines’: absent/ present,

• X [214] ‘[DFC 127] Anal fin spine’: absent/ present,

• X [215] ‘[DFC 128] Paired fin spines’: absent/ present,

• T [216] ‘[DFC 129] Median fin spine insertion’: ‘shallow, not greatly deeper than dermal bones/scales’/ deep,

• X [217] ‘[DFC 130] Intermediate fin spines’: absent/ present,

• T [218] ‘Fin spine cross-section’: ‘Round or horseshoe shaped’/ ‘Flat-sided, with rectangular profile’,

• T [219] ‘Intermediate spines when present’: ‘one pair’/ ‘multiple pairs’,

• N [220] ‘[DFC 131] Prepectoral fin spines’: ‘absent’/ present,

• N [221] ‘[DFC 132] Fin spines with ridges’: absent/ present,

• N [222] ‘[DFC 133] Fin spines with nodes’: absent/ present,

• N [223] ‘[DFC 134] Fin spines with rows of large retrorse denticles’: absent/ present,

• N [224] ‘Expanded spine rib on leading edge of spine’: absent/ present,

• T [225] ‘Spine ridges’: ‘converging at the distal apex of the spine’/ ‘converging on leading edge of spine’,

• X [226] ‘[DFC 135] Synarcual’: absent/ present,

• X [227] ‘Series of thoracic supraneurals’: absent/ present,

• T [228] ‘[DFC 136] Number of dorsal fins, if present’: one/ two,

• T [229] ‘Posterior dorsal fin shape’: ‘base approximately as broad as tall, not broader than all of other median fins’/ ‘base much longer than the height of the fin, substantially longer than any of the other dorsal fins’,

• N [230] ‘Basal plate in dorsal fin’: absent/ present,

• T [231] ‘Branching radial structure articulating with dorsal fin basal plate’: absent/ present,

• X [232] ‘[DFC 137] Anal fin’: absent/ present,

• N [233] ‘Basal plate in anal fin’: absent/ present,

• T [234] ‘[DFC 138] Caudal radials’: ‘extend beyond level of body wall and deep into hypochordal lobe’/ ‘radials restricted to axial lobe’,

• X [235] ‘Supraneurals in axial lobe of caudal fin’: absent/ present,

• N [236] ‘Epichordal lepidotrichia in caudal fin’: absent/ present,

43 taxa, 137 characters, of which 60 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 13, 14, 15, 36, 52, 54, 55, 57, 60, 61, 62, 65, 72, 76, 78, 79, 80, 81, 82, 84, 89, 91, 92, 106, 107, 108, 110, 111, 113, 123, 124, 125, 128

• Neomorphic: 4, 6, 68, 71, 73, 74, 77, 87, 88, 93, 94, 95, 98, 112, 117, 122, 126, 129, 130, 131, 132, 133, 134, 135, 136 ; with state 1 denoting absence: 120, 121

Details:

• X [1] Femoral_trichobothria / absent present,

• X [2] Tarsal_organ / exposed capsulate,

• X [3] Tarsal_trichobothria / absent present,

• N [4] Tarsal_trichobothrial_rows / l ‘2+’, Neomorphic as additional rows seen as a neomorphism

• X [5] Metatarsal_trichobothria / ‘1-2’ ‘>2’,

• N [6] Tarsal_trichobolhria / normal longer_distally, ‘normal’ suggests that this is interpreted as the ancestral condition

• X [7] Palpal_tibial_trichobothria / absent present,

• X [8] Trichobothrial_base / smooth trans._ridge long._ridge,

• X [9] Trichobothrial_base / simple notched,

• X [10] Trochanteral_notch / absent present,

• X [11] Tarsal_claws / 3. 2.,

• X [12] Claw_tufts / absent present,

• T [13] Calamistral_rows / 2. l. 3.,

• T [14] Calamistrum / linear oval,

• T [15] Calamistrum_origin / basal median,

• X [16] Male_palpal_femoral_thoms / absent present,

• X [17] Female_palpal_femoral_thoms / absent present,

• X [18] Feathery_hairs / absent present,

• X [19] Hair / plumose pseudoserrate serrate,

• X [20] Metatarsal_preening_combs / absent present,

• X [21] Tibial_ventral_spine_number / ‘<7’ ‘>7’,

• X [22] Male_metatarsus_1 / unmodified modified,

• X [23] Male_tibial_crack / absent present,

• X [24] Serrate_accessory_claw_setae / absent present,

• X [25] Fernale_posterior_leg_scopula / absent present,

• X [26] Deinopoid_tarsal_comb / absent present,

• X [27] Carapace_shape / oval square round,

• X [28] Clypeal_hood / absent present,

• X [29] Thickened_setae_at_fang_base / absent present,

• X [30] Male_chelicerae / normal bowed,

• X [31] Chelicerae / normal small,

• X [32] Chelicerae / free fused_at_base,

• X [33] Cheliceral_teeth / present absent,

• X [34] Cheliceral_chela / absent present,

• X [35] Cheliceral_boss / absent present,

• T [36] Cheliceral_bow / small large,

• X [37] Chilum / absent median bilateral,

• X [38] Tapetum / primitive canoe grate absent,

• X [39] Posterior_eye_row / straight recurved,

• X [40] Serrula_tooth_rows / multiple single,

• X [41] Sigilla / present absent,

• X [42] Medial_cheliceral_concavity / present absent,

• X [43] ‘Venom, gland’ / cheliceral carapace absent,

• X [44] Coxal_gland_duct / convoluted simple,

• X [45] 5th_endostemite_invagination / present absent,

• X [46] Midgut_cheliceral_diverticula / present absent,

• X [47] Pharynx_musele_origin / carapace rostrum,

• X [48] Intestine / ‘M-shaped’ straight,

• X [49] Heart_ostia_nurnber / 4. 3_or_2,

• X [50] Anal_tubercle / small large,

• X [51] Posterior_spiracIes / 2. l.,

• T [52] Posterior_spiracies / wide narrow,

• X [53] Posterior_respiratory_system / booklungs tracheae,

• T [54] Median_tracheae / simple branched,

• T [55] Lateral_tracheae / simple branched absent,

• X [56] Epiandrous_spigots / present absent,

• T [57] Epiandrous_spigots / dispersed 2_bunches,

• X [58] Spinn._cuticle / armul. ridged squam. papill.,

• X [59] Cribellum / present absent,

• T [60] Cribellum / entire divided,

• T [61] Cribellate_spigots / uniform clumped,

• T [62] Cribellate_spigots / strobilate claviform,

• X [63] Tartipores / absent present,

• X [64] ALS_field_margin / narrow broad,

• T [65] ALS_MAP / clustered dispersed,

• X [66] ALS_MAP_number / ‘>3’ 3. 2. l.,

• X [67] ALS_MAP_nubbin / absent present,

• N [68] ALS_MAP_nubbin / one extra, [‘Extra’ interpreted as neomorphism]

• X [69] ALS_piriform_margin / rounded flat sharp,

• X [70] PMS_mAP / absent present,

• N [71] PMS_mAP_number / l. ‘>l’,

• T [72] PMS_mAP_position / medianlanterior posterior,

• N [73] PMS_mAP_nubbins / absent present,

• N [74] PMS_mAP_nubbins / l. 2.,

• X [75] PMS_aciniform / present absent,

• T [76] PMS_aciniform_number / ‘>3’ ‘1-3’,

• N [77] PMS_paracribellars / absent present,

• T [78] PMS_paracribellar / at_apex ‘mid-field’,

• T [79] PMS_paracribellar / bunched encircling,

• T [80] PMS_paracribellar_shafts / single_bases grouped,

• T [81] PMS_paracribellar_bases / round long narrow,

• T [82] PMS_paracribellar / deinopoid strobilate floppy,

• X [83] PMS_cylindrical_spigots / absent present,

• T [84] PMS_cylindrical_spigot_number / ‘1-2’ many,

• X [85] PLS_cuticular_finger / absent present,

• X [86] PLS_aggregate_gland_spigot / absent present,

• N [87] PLS_paracribellar / absent present,

• N [88] PLS_paracribellar_number / l. ‘2-3’,

• T [89] PLS_paracribellar / separate ‘+ modified spigot’,

• X [90] PLS_modified_spigot / absent present,

• T [91] PLS_modified_spigot / in_field segregated,

• T [92] PLS_apical_segment / domed conical elongate,

• N [93] PLS_apical_enlarged_seta / absent present,

• N [94] Tibial_retroapical_process / absent present,

• N [95] Tibial_retroapical_process / simple complex,

• X [96] Tibial_ventroapical_process / absent present,

• X [97] Tibial_dorsal_process / absent apical proximal,

• N [98] Tibial_dorsal_process / simple complex_folded,

• X [99] Tibial_prolateral_process / absent present,

• X [100] Paracymbium / absent present,

• X [101] Cymbial_dorsal_scopula / absent present,

• X [102] Pyriform_bulb / absent present,

• X [103] Tegular_locking_lobe / absent present,

• X [104] Subtegular_loeking_lobe / absent present,

• X [105] Conductor / present absent tegular_groove,

• T [106] Conductor / sclerotized hyaline,

• T [107] Scl._C / embr._E opp._E cent. Ulo Oe Th,

• T [108] C_embr._E / apex_only desid dict met apical,

• X [109] Median_apophysis / present absent,

• T [110] Median_apophysis / convex concave,

• T [111] Median_apophysis / fixed flexibly_attached,

• N [112] Extra_tegular_processes / absent present,

• T [113] Extra_tegular_process / conical STP membr.,

• X [114] Palpal_tarsus_M29_muscle / present absent,

• X [115] Female_genitalia / haplogyne entelegyne,

• X [116] Epigynum / absent present,

• N [117] Epigynum_teeth / absent present,

• X [118] Convoluted_vulva / absent present,

• X [119] Posterior_receptaculum / absent present,

• I [120] Cribellate_silk_axial_lines / present absent,

• I [121] Cribellate_silk_reserve_arp / present absent,

• N [122] Cribellate_silk_nodules / absent present,

• T [123] Cribellate_fibrils / cylindrical flat,

• T [124] Cribellate_silk / uniform puffed,

• T [125] Web_posture / inverted erect,

• N [126] Orb_web_architecture / absent present,

• X [127] Deinopid_web_architecture / absent present,

• T [128] Combing_leg_support / fixed_L3 mobile_L4,

• N [129] Frame_construction / absent present,

• N [130] Radius_construction / absent ‘CR-ed’ doubled,

• N [131] Hub_construction / absent present,

• N [132] Temporary_spiral_construction / absent present,

• N [133] Sticky_spiral_construction / absent present,

• N [134] Sticky_silk_iocalization / absent oLl iL4,

• N [135] L4_sticky_silk_shift / absent present,

• N [136] ‘Non-sticky-sticky line grip’ / otherwise w_L4,

• X [137] Wrap_attack / absent present,

68 taxa, 308 characters, of which 97 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 2, 4, 48, 54, 55, 62, 65, 72, 73, 74, 78, 80, 81, 96, 113, 115, 116, 120, 122, 138, 139, 140, 141, 142, 147, 148, 149, 151, 152, 161, 162, 163, 170, 173, 178, 181, 182, 183, 186, 187, 188, 189, 190, 200, 204, 208, 213, 215, 216, 217, 218, 219, 220, 221, 222, 224, 235, 242, 246, 248, 252, 258, 270, 271, 272, 273, 275, 276, 277, 278, 279, 280, 281, 282, 283, 287, 288, 291, 295, 297, 305

• Neomorphic: 17, 42, 75, 79, 121, 150, 175, 223, 234, 236, 237, 274 ; with state 1 denoting absence: 144, 191, 233, 269

Details:

• N [42] Hp22, Neomorphic as ‘ordered’

56 taxa, 104 characters, of which 66 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 1, 2, 3, 5, 9, 15, 19, 23, 29, 32, 34, 35, 36, 37, 38, 39, 43, 44, 45, 46, 47, 48, 49, 51, 53, 54, 55, 56, 57, 58, 59, 62, 63, 66, 67, 68, 70, 71, 72, 73, 74, 76, 77, 78, 79, 83, 86, 89

• Neomorphic: 30, 31, 33, 40, 41, 42, 50, 52, 61, 64, 65, 69, 75, 80, 81, 82, 85, 88

Details:

• T [1] Habit / ‘shrub (n)’ tree herb,

• T [2] Stem_vasculature / circular cortical scattered,

• T [3] Cambial_initials / long short,

• X [4] Xylem_conduction / tracheids ‘vessels foraminate perforate (n’ vessels_scalariform_perforate vessels_simple_perforate,

• T [5] Vessel_distribution / whole_plant roots_only,

• X [6] Primary_pitting / late_ontogeny_of_tracheary_elem early_ontogeny_of_tracheary_ele,

• X [7] ‘Sieve-tube element companion ce’ / absent present,

• X [8] Secondary_phloem / ‘non-stratified’ tangentially_stratified,

• T [9] ‘Sieve-tube plastic inclusions’ / starch protein,

• X [10] Pith_structure / nonseptate sclereid_septate,

• X [11] Sclereids / ‘non-hippocrepiform’ hippocrepiform,

• X [12] Corticalsclerenchyma_sheath / absent present,

• X [13] Laticifers / absent present,

• X [14] Oil_cells / absent present,

• T [15] Nodal_anatomy / ‘unilacunar two-trace’ ‘unilacunar one-trace’ trilacunar multilacunar,

• X [16] Nodal_morphology / ‘non-swollen’ swollen,

• X [17] Axillary_buds / absent present multiple,

• X [18] Phyllotaxy / alternate opposite,

• T [19] Ptyxis / circinate adplicate conduplicate supervolute involute,

• X [20] Leaf_type / compound simpl,

• X [21] Leaf_peltate / absent present,

• X [22] Leaf_base / simple ‘sheathing (n)’ stipulate,

• T [23] Leaf_venation / dichotomous pinnate palmate,

• X [24] Leaf_margin / entire toothed,

• X [25] Leaf_persistence / evergreen deciduous,

• X [26] Leaf_pubescence / absent present,

• X [27] Foliar_astrosclereids / absent present,

• X [28] Foliar_cystoliths / absent present,

• T [29] Stomates / anomocytic paracytic laterocytic ‘stephanocytic (n)’ ‘encyclocytic (n)’,

• N [30] Infloresence / ‘absent (flowers solitary)’ present,

• N [31] Floral_pedicel / present absent,

• T [32] Floral_primordium_symmetry / radial ‘bilateral (dorsiventral)’,

• N [33] Calyptra / absent present,

• T [34] Perianth_arrangement / helical cyclic_and_trimerous cyclic_and_four_or_five_merous absent,

• T [35] Petals / absent tepaloid staminoidal,

• T [36] Perianth_androecium_alignment / alternate opposite,

• T [37] Sexual_expression / dioecious monoecious monoclinous,

• T [38] Stamen_type / connective_with_marginal_sporan ‘laminar (with adaxial or abaxia’ ‘filamental (with terminal spora’,

• T [39] Stamen_arrangement / helical cyclic,

• N [40] Staminal_osmophore / absent present,

• N [41] Monadelphy / absent present,

• N [42] Filament_appendage / absent present,

• T [43] Anther_dehiscence / longitudinal bifurcate valvate,

• T [44] Connective_hypodermic / absent ‘endothecial-like’,

• T [45] Tapetum / glandular amoeboid,

• T [46] Microsporogenesis / simultaneous successive,

• T [47] Pollen_unit / monads tetrads,

• T [48] Pollen_size / ‘medium (20-50 pm)’ ‘large (greater than 50pm)’ ‘small (less than 20pm)’,

• T [49] Pollen_shape / ‘boat-shaped’ intermediate globose,

• N [50] Pollen_saccate / absent present,

• T [51] Pollen_apertures / monosulcate ‘disulcate (n)’ tricolpate ‘holoaperturate (n)’ ‘ulcerate (n)’ ‘polyaperturate (n)’,

• N [52] Apertures / free fused,

• T [53] Examine_sculpture / psilate ‘punctate (n)’ reticulate ‘striate (n)’,

• T [54] Pollen_exine_surface / smooth spinulose spinose,

• T [55] Exine_structure / alveolar granular ‘solid (n)’ columellate ‘chanellate (n)’,

• T [56] Endexine / continuous discontinuous absent,

• T [57] Pollen_tube_growth / suspended penetrating,

• T [58] Microgametophyte / prothallial_and_sterile_cells_p sterile_cells_absent prothallial_and_sterile_cells_a,

• T [59] Microgametophyte_final_mitotic / ‘post-pollination’ prepollination,

• X [60] Sperm / flagellated ‘non-flagellated’,

• N [61] Inner_staminodes / present absent,

• T [62] Carpel_type / open closed,

• T [63] Carpel_arrangement / helical ‘cyclic (n)’ solitary,

• N [64] Floral_cup / absent present,

• N [65] Gynoecial_nectaries / absent present,

• T [66] Placentation / marginal laminar apical basal parietal exile,

• T [67] Ovule_type / orthotropous anatropous,

• T [68] Ovule_number / uniovulate_or_biovulate multiovulate,

• N [69] Micropylar_tube / absent present,

• T [70] Integuments / unitegmic bitegmic,

• T [71] Nucellar_cuticle / thin thick,

• T [72] Megagametophyte_meiosis / monosporic tetrasporic,

• T [73] Megaspore_wall / thick thin,

• T [74] Megagametophyte_apex / cellular ‘free-nuclear’,

• N [75] Fertilization / single double,

• T [76] Fruit_type / dehiscent berry achene drupe,

• T [77] Seed_size / ‘large (>5 mm)’ ‘small (<5 mm)’,

• T [78] Seed_coat_anatomy / endotestal mesotestal exotestal tegmic,

• T [79] Seed_aril / exarillate arillate,

• N [80] Sarcotestum / absent present,

• N [81] Seed_chalazal_wings / absent present,

• N [82] Seeds_operculate / absent present,

• T [83] Endosperm_development / cellular nuclear helobial,

• [84] Endosperm_type / absent present perisperm,

• N [85] Ruminate_endosperm / absent present,

• T [86] Embryogenesis / ‘free-nuclear’ onagrad_type asterad_type caryophyllad_type solanum_type,

• X [87] Embryo_size / minute large,

• N [88] Embryo_with_feeder / absent present,

• T [89] Germination / hypogeal epigeal,

• X [90] Root_system / allorhizic homorhizic,

• X [91] Cotyledon_number / two two_laterally_fused one ‘three or more (n)’,

• X [92] Lignin_subunits / vanillin syringal_aldehyde,

• X [93] Hydrolyzable_tannin / absent present,

• X [94] Proanthocyanidin_leucoanthocyan / present absent,

• X [95] Cyanidin / absent present,

• X [96] ‘Methoxy-anthocyanidin’ / absent present,

• X [97] ’3" 4" 5“-Methoxy-flavonol myric’ / absent present,

• X [98] ’7 3" 4" or 5“-Methoxy-Ravonol’ / absent present,

• X [99] ‘8-Oxygenated-flavonol’ / absent present,

• X [100] Dihydrofiavonol / absent present,

• X [101] Apigenin / absent present,

• X [102] ‘C-Glycosylfavone’ / absent present,

• X [103] Isoflavone / absent present,

• X [104] Benzylisoquinoline / absent present,

20 taxa, 93 characters, of which 22 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 2, 3, 5, 6, 12, 16, 18, 20, 24, 26, 28, 29, 65, 70, 80

• Neomorphic: 13, 23, 25, 41, 55, 57, 64

Details:

• X [1] 1 Frontals and parietals flat dorsally (0) or frontals and parietals thickened and vaulted to form a cranial dome (1) (Sereno, 2000: character 4).

• T [2] 2 Frontoparietal dome with discrete orbital and postorbital lobes (0) or orbital and postorbital lobes confluent (1).

• T [3] 3 Frontoparietal dome short and broad (0) or frontoparietal dome elongate, dome roughly 150% as long as wide (1).

• X [4] 4 Frontals free (0) or interfrontal suture and frontoparietal sutures closed (1).

• T [5] 5 Frontal, orbital processes distinct from dome (0) or completely incorporated into dome (1).

• T [6] 6 Frontals, dome narrow over orbits, (0) or dome over orbits transversely expanded (1).

• X [7] 7 Frontal, nasal processes with a flat dorsal surface (0) or nasal processes convex, forming a low nasal boss (1) or nasal processes elevated above prefrontals to create a tall nasal boss (2) (modified from Williamson and Carr, 2002: character 42).

• X [8] 8 Frontal, nasal process smooth or bearing low nodes (0) or nasal process bearing enlarged nodes (1).

• X [9] 9 Frontal, nasal processes elongate, about 25% length of frontoparietals (0) or shortened, less than 25% length of frontoparietals (1) highly reduced, 15% or less length of frontoparietals (2).

• X [10] 10 Frontal contributes to rim of orbit (0) or frontal excluded from orbit by supraorbital ossifications (1) (Sereno, 2000: character 8).

• X [11] 11 Frontal lacks articulation with accessory orbital ossifications, medial wall of orbit open (0) or frontal articulates with accessory orbital ossifications (1).

• T [12] 12 Frontal with large articular surface for supraorbital 1 (0) or reduced articulation between frontal and supraorbital 1 (1).

• N [13] 13 Frontal, orbital roof with large vascular foramena and associated blood vessel grooves: absent (0) or present (1).

• X [14] 14 Frontals subequal to, or longer than, parietals in length (0) or parietals substantially longer than frontals (1).

• X [15] 15 Parietals, articulation with postorbitals limited or absent (0) or extensive (1) (Sereno, 2000: character 14).

• T [16] 16 Parietals, smooth or bearing small tubercles (0) or bearing large nodes on caudal margin (1).

• X [17] 17 Parietals, covered dorsally by supratemporal fossae (0), or parietals overhang or entirely roof over adductor chamber (1) (modified from Williamson and Carr, 2002: Character 40).

• T [18] 18 Parietals, dorsal roof of adductor chamber shallow (0) or deeply excavating dome posterodorsally (1).

• X [19] 19 Parietals: broadly participate in caudal margin of occiput (0), largely excluded by squamosals (Williamson and Carr, 2002: character 44).

• T [20] 20 Parietals, dome only incorporates anterior part of parietals (0), or dome fully incorporates parietal, eliminating parietosquamosal shelf (1) (Williamson and Carr, 2002: character 51) or dome caudally extended, overhanging parietosquamosal shelf (2).

• X [21] 21 Parietal, postorbital and squamosal form margins of supratemporal fenestrae (0) or supratemporal fenestrae closed by contact between parietals, squamosals, and postorbitals (1).

• X [22] 22 Squamosal, enlarged squamosal nodes absent (0) or present (1)

• N [23] .. or hypertrophied (2) ORDERED (modified from Sereno, 2000: character 16; Williamson and Carr, 2002: character 39).

• T [24] 23 Squamosal, nodes conical or button-shaped (0) or developed as spikelike horns (1).

• N [25] 24 Squamosal, squamosal nodes in a single row (0), or single row with distinct corner node below (sometimes with 1-2 small accessory nodes) (1), 2 or more rows (2) (Williamson and Carr, 2002: characters 36, 37) UNORDERED

• T [26] 25 Squamosal, enlarged nodes in row on caudal margin number 5 or more (0), 3-4 (1) (Sullivan, 2003: character 42).

• X [27] 26 Squamosals straight in caudal view (0) or tips of squamosals pendant in caudal view (1) (Williamson and Carr, 2002: 37).

• T [28] 27 Squamosal, Parietosquamosal shelf deep laterally (0), or thins laterally in caudal view (1) (Modified from Williamson and Carr, 2002).

• T [29] 28 Squamosal contribution to dome limited (0) or squamosal extends dorsally to the same extend as the postorbital (1).

• X [30] 29 Squamosal, with restricted exposure on occiput (0) or with broad exposure on occiput (1) (Sereno, 2000: character 7).

• X [31] 30 Squamosal, prominent crescentic scars on occipital surface absent (0) or present (1).

• X [32] 31 Squamosal, posterolateral corner massive and projecting caudally to form a pedicel: absent (0) or present (1).

• X [33] 32 Squamosals separated by parietal (0) or squamosals contact posteriorly (1).

• X [34] 33 Caudal margins of parietals and squamosals dorsal to occiput (0), project caudally beyond occiput forming a parietosquamosal shelf (1) (Sereno 2000, character 34).

• X [35] 34 Postorbital, postorbital nodes small or absent (0) or bearing a distinct row of enlarged nodes along lateral margin (1).

• X [36] 35 Postorbital with distinct postorbital node row descending postorbital bar: absent (0) or present (1) .

• X [37] 36 Postorbital-jugal bar slender (0) or robust (1) (Sereno, 2000: character 15).

• X [38] 37 Postorbital and squamosal form a slender bar (0) or a robust shelf, sometimes incorporated into dome (1).

• X [39] 38 Postorbital-squamosal, lateral margin straight in dorsal view (0) or strongly convex in dorsal view, with squamosal and postorbital forming a crown around the skull (1).

• X [40] 39 Nasals, nodes on nasals absent (0), present (1)

• N [41] .. , hypertrophied (2) ORDERED (Williamson and Carr, 2002: character 52).

• X [42] 40 Nasals flat dorsally, resulting in a dorsally flat rostrum (0), or nasals arched transversely resulting in a convex dorsal surface of the rostrim (1).

• X [43] 41 Nasals short, no more than 150% as long as wide, rostrum short (0), snout elongate, paired nasals roughly twice as long as wide, rostrum elongate (1) (modified from Sereno, 2000, character 39).

• X [44] 42 Supraoccipital: occipital depression above foramen magnum absent (0) or present (1) (Williamson and Carr, 2000, character 45).

• X [45] 43 Supraoccipital: epaxial muscle scars: restricted to exoccipital (0) or extend medially onto supraoccipital (1) (Williamson and Carr, 2002: character 47).

• X [46] 44 Basioccipital: basal tubera massive (0) or thin and plate-like (1) (Sereno, 2000: character 18).

• X [47] 45 Basioccipital: occipital condyle weakly differentiated from body of basioccipital (0), or occipital condyle borne on an elongate, pillar-like neck (1)

• X [48] 46 Lateral temporal fenestra ventral margin elongate (0) or anteroposteriorly short (1).

• X [49] 47 Jugal articulates with quadrate: absent (0) or present (1) (Sereno, 2000: character 37).

• X [50] 48 Quadrate articulates high on skull (0) or quadrate articulates low on descending process of squamosal (1).

• X [51] 49 Quadrate shaft with a shallow fossa largely covering the lateral surface (0) or quadrate lateral fossa reduced (1).

• X [52] 50 Premaxilla: external nares retracted from anteroventral margin of premaxilla (0) or nares extend down onto anteroventral surface of premaxilla (1).

• X [53] 51 Premaxillae: ventral margin of premaxillae with sharp-edged tomial surface (0) or broad and flattened lip-like rugosity (1).

• X [54] 52 Premaxillae, maxillary ramus strongly arched to form a distinct premaxilla-maxilla notch: absent (0) or present (1) (modified from Sereno, 2000: character 5).

• N [55] 53 Prootic-basisphenoid plate: absent (0) or present (1) (Sereno, 2000, character 27).

• X [56] 54 Pterygoquadrate rami, posterior projection of ventral margin: weak (0); pronounced (1) (Sereno, 2000; character 26).

• N [57] 55 Exoccipital, scars for attachment of epaxial muscles on paroccipital processes: absent or indistinct (0); prominent (1) (Williamson and Carr, 2002; character 47).

• X [58] 56 Exoccipital: exoccipital laterally projecting (0) strongly pendant (1).

• X [59] 57 Dentary forms anterior margin of coronoid process (0) or coronoid forms anterior margin of coronoid process (1).

• X [60] 58 Dentary anterior end deep in lateral view (0) or anterior end of dentary shallow and strongly tapered in lateral view (1).

• X [61] 59 Coronoid process of mandible tall and subtriangular (0) or developed as a low, crescentic flange (1).

• X [62] 60 Surangular and angular, lateral fossa present (0) or absent (1).

• X [63] 61 Skull, external surfaces of elements of the skull roof, face, and mandible: smooth and unornamented (0) or rugose, ornamented with nodes, tubercles, and pits of varying size (1).

• N [64] 62 Dentition, premaxillary and anterior dentary teeth lack denticles (0) or denticles present (1).

• T [65] 63 Dentition, premaxillary and anterior dentary teeth subconical (0) laterally compressed and bladelike (1).

• X [66] 64 Cervical vertebrae, ventral surface of centrum rounded (0) or keeled (1).

• X [67] 65 Vertebrae, zygapophyseal articular surfaces planar (0) or with ridge-and-groove structure (1).

• X [68] 66 Sacral ribs subrectangular (0) or elongate and strap shaped (1).

• X [69] 67 Proximal caudal vertebrae, neural spines posteriorly inclined (0) or projecting dorsally (1).

• T [70] 68 Tail, ossified tendons rodlike(0) or massive and fusiform (1) (Sereno, 2000: character 20).

• X [71] 69 Scapular blade broad (0) or narrow and strap-shaped (1) (Sereno, 2000: character 2).

• X [72] 70 Sternum plate-like (0) or shafted (1) (Sereno, 2000: character 21).

• X [73] 71 Humerus elongate (0) or short, less than 50% length of femur (1) ( Sereno, 2000: character 10).

• X [74] 72 Humerus, deltopectoral crest prominent (0) or reduced (1) (Sereno, 2000: character 12).

• X [75] 73 humeral shaft straight (0) or bowed (1) (Sereno, 20: character 11).

• X [76] 74 Ulna robust (0) or more slender than radius (1).

• X [77] 75 Ilium, postacetabular wing with strong medial curvature, absent (0) or present (1) (Sereno, 2000).

• X [78] 76 Ilium, prominent medial process absent (0) or present (1) (Sereno, 2000) or greatly expanded, extending 2/3 length of postacetabular process (2).

• X [79] 77 Ilium, anterior process a vertical blade (0) or transversely expanded (1)

• T [80] .. or strongly expanded, spatulate (2) (modified from Sereno, 2000: Character 3) ORDERED.

• X [81] 78 Ilium, dorsal margin with distinct dorsal shelf: absent (0) or present (1).

• X [82] 79 Ilium, anterior blade with little or no lateral deflection (0) or strong lateral deflection, preacetabular process bent laterally by 30 degrees (1) (Sereno, 2000: character 22).

• X [83] 80 Ilium, medial tab (???) absent or present (Sereno, 2000; character 23).

• X [84] 81 Ilium: pubic peduncle mediolaterally narrow (0) or strongly transversely expanded and dorsoventrally compressed (1) (2000: character 30).

• X [85] 82 Ilium posterior wing straight in lateral view (0) or strongly curved downwards (1).

• X [86] 83 Ilium, postacetabular blade short (0) or elongate, longer than preacetabular blade (1).

• X [87] 84 Pubis, contribution to acetabulum extensive (0), or reduced (1) (modified from Sereno, 2000: character 31).

• X [88] 85 Pubis, prepubic process distally expanded: absent (0) or present (1).

• X [89] 86 ischium straight or weakly curved (0) or ischium strongly bent, angle of at least 30º between proximal and distal ends of shaft (1).

• X [90] 87 Femur, femoral fourth trochanter elongate and pendant (0) or developed as a low flange (1).

• X [91] 88 Femur, femoral head weakly separated from greater trochanter (0) or femoral head and greater trochanter separated by a deep notch in anterior or posterior view (1).

• X [92] 89 Femur, lesser trochanter proximodistally short, arises from proximal 25% of femoral shaft (0), or lesser trochanter extended distally, arises from proximal 33% of femoral shaft.

• X [93] 90 Fibular mid-shaft diameter greater than 20% tibia diameter (0) or no more than 20% diameter of tibia (1) (Sereno, 2000: character 13).

40 taxa, 150 characters, of which 67 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 34, 35, 49, 50, 53, 56, 57, 58, 59, 60, 62, 69, 70, 78, 84, 86, 88, 93, 122, 124, 125, 126, 128, 130, 131, 132, 133

• Neomorphic: 13, 14, 15, 16, 17, 18, 19, 20, 52, 54, 55, 63, 64, 65, 68, 71, 72, 74, 75, 79, 80, 81, 82, 83, 85, 87, 89, 90, 91, 92, 120, 121, 127, 134 ; with state 1 denoting absence: 61, 66, 67, 73, 123, 129

Details:

• X [1] 001. Subarcuate fossa.—-Present (0); absent (1) (Novacek, 1986).

• X [2] 002. Shape of tegmen tympani (ordered).—-Uninflated, forms thin lamina lateral to facial nerve canal (0); inflated, forms barrel-shaped ossi?cation lateral to the facial nerve canal (1);

• X [3] …. hyperinflated, transverse width greater than or equal to width of promontorium (2) (modi?ed from Cifelli, 1982; Geisler and Luo, 1998).

• X [4] 003. Anterior process of petrosal.—-Absent (0); present, anterior edge of tegmen tympani extending far ante- rior to edge of pars cochlearis (1) (Luo and Marsh, 1996; Geisler and Luo, 1998).

• X [5] 004. Fossa fortensortympanimuscle.—-Shallow, anteroposteriorly elongate fossa (0); circular pit, no groove (1); circular pit with deep tubular anterior groove (2); long narrow groove between tegmen tympani and promontorium (3) (Luo and Marsh, 1996; Geisler and Luo, 1998).

• X [6] 005. Transpromontorial sulcus for internal carotid artery.—- Present, forms anteroposterior groove on promontorium, medial to fenestrae rotunda and ovalis (0); absent (1) (Cifelli, 1982; Thewissen and Domning, 1992).

• X [7] 006. Sulcus on promontoriumfor proximalstapedial artery.—- Present, forms a groove that branches from the transpromontorial sulcus anteromedial to the fen- estra rotunda and extends to the medial edge of the fenestra vestibuli(0); absent (1) (Cifelli, 1982; Wible, 1987; Thewissen and Domning, 1992).

• X [8] 007. Mastoid process of petrosal.—-Exposed externally on posterior face of braincase as a triangle between lambdoidal crest of the squamosal dorsolaterally, the exoccipital ventrally, and the supraoccipital me- dially (0); not exposed posteriorly, lambdoidal crest of squamosal in continuous contact with exoccipital and supraoccipital (1).

• X [9] 008. Stylomastoid foramen.—-Complete, ectotympanic contacts tympanohyal laterally and petrosal me- dially, in some cases ectotympanic separated from petrosal by a narrow (< 1 mm) ?ssure (0); incom- plete, ectotympanic does not contact the petrosal either anterior or posterior to the fenestra rotunda, medial side of foramen open (1) (modi?ed from Geisler and Luo, 1998).

• X [10] 009. Facial nerve sulcus distal to stylomastoid foramen.—- Absent (0); present, anterior wall of sulcus formed by squamosal (1); present, anterior wall formed by mastoid process of petrosal(2); anterior wallformed by meatal tube of ectotympanic (3) (modi?ed from Geisler and Luo, 1996, 1998). Some taxa coded as polymorphic for this character are not truly poly- morphic, and the character state is simply ambigu- ous, but certain states can be eliminated from the list of possibilities.

• X [11] 010. Articulation of pars cochlearis with basisphenoid/basioc- cipital.—-Present (0); absent (1) (Thewissen and Domning, 1992).

• X [12] 011. Ectotympanic.—-Simple ring, no bulla formation (0); medial edge expanded into bulla (1) (derived from Novacek, 1977; MacPhee, 1981; Geisler and Luo, 1998).

• N [13] 012. Pachyosteosclerotic involucrum of bulla.—-Absent (0); present (1) (Thewissen, 1994).

• N [14] 013. Lateral furrow of tympanic bulla.—-Absent(0);present, forms a groove on the lateral surface of the ectotympanic bulla anterior to the base of the sigmoid process or meatal tube (1) (Geisler and Luo, 1998).

• N [15] 014. Median furrow of tympanic bulla for tympanohyal (ordered).—-Absent (0); median notch on posterior rim of bulla (1);

• N [16] …. prominent anteroposteriorly ori- ented furrow splits the ventral surface of the bulla into medial and lateral halves (2) (Geisler and Luo, 1998).

• N [17] 015. Articulation of ectotympanic bulla to squamosal (ordered).—-Broad articulation with medial base of postglenoid process (0); circular facet on short entoglenoid process (1);

• N [18] …. contact with a crest of the entoglenoid process (2);

• N [19] …. contact absent(3)(Geisler and Luo, 1998).

• N [20] 016. Contact between exoccipital and ectotympanic bulla.—- Absent (0); present (1) (Geisler and Luo, 1998).

• X [21] 017. Sigmoid process(= anterior crus of tympanic ring).—- Absent (0); present, forms transverse plate that projects laterally from the anterior crus of the ectotympanic ring and forms the anterior wall of the external auditory meatus (1) (modi?ed from Thewis- sen, 1994; Geisler and Luo, 1998).

• X [22] 018. Sigmoid process shape.—-Thin and transverse plate (0); broad and flaring, base of the sigmoid process forms dorsoventral ridge on lateral surface of ectotympanic bulla (1) (Geisler and Luo, 1998).

• X [23] 019. Ectotympanic part of the meatal tube (ordered).—- Absent (0); present but short, length of tube < 30% maximum width of bulla (1);

• X [24] …. present and long, length > 60% maximum width of bulla (2) (Geisler and Luo, 1998).

• X [25] 020. Squamosal portion of external auditory meatus.—- Absent or shallow, depth < 25% of transverse length (0); deep groove, depth > 35% of transverse length (1).

• X [26] 021. Basioccipital crests (falcate processes).—-Absent (0); present, forming ventrolaterally flaring basioccipital processes (1) (derived from Barnes, 1984; modi- ?ed from Thewissen, 1994).

• X [27] 022. Internal carotid foramen.—-Absent or confluent with the piriform fenestra (0); present at basisphe- noid/basioccipital suture with lateral wall of fora- men formed by both these bones and thus sepa- rated from the piriform fenestra (1) (Geisler and Luo, 1998).

• X [28] 023. Postglenoid foramen (ordered).—-Present, enclosed by squamosal, immediately posterior to post- glenoid process (0); present, medial to pet- rosal/squamosal suture (if bulla present, a sec- ondary ventral opening between bulla and squamosal may form) (1);

• X [29] …. absent (2) (Novacek, 1986; Geisler and Luo, 1998).

• X [30] 024. Foramen for ramus superior of stapedial artery.—- Present (0); absent (1) (modi?ed from Novacek, 1986; Thewissen and Domning, 1992).

• X [31] 025. Position of foramen for ramus superior of stapedial artery.—-In petrosal/squamosal suture on dorsolat- eral edge of epitympanic recesss (0); anterolateral to epitympanic recess (1) (modi?ed from Geisler and Luo, 1998). Cannotbescored fortaxa thatlack a fora- men for the ramus superior of the stapedial artery.

• X [32] 026. Foramen ovale.—-Anterior to glenoid fossa, posterior wall formed by alisphenoid (0); medial to glenoid fossa, posterior wall formed by alisphenoid (1); me- dial to glenoid fossa, posterior wall formed by pet- rosal (2); posterior to glenoid fossa (3).

• X [33] 027. Alisphenoid canal (alar canal).—-Absent (0); present (1) (Novacek, 1986; Thewissen and Domning, 1992).

• T [34] 028. Anterior opening of alisphenoid canal.—-Within sphe- norbital ?ssure, cannot be distinguished from sphe- norbital ?ssure in lateral view (0); posterior to sphe- norbital ?ssure (1) (Novacek, 1982; Thewissen and Domning, 1992).

• T [35] 029. Posterior opening of alisphenoid canal.—-Wellseparated from foramen ovale (0); within a deep groove or recess with the foramen ovale (1) (de- rived from Zhou et al., 1995).

• X [36] 030. Foramen rotundum.—-Absent, maxillary division of trigeminal nerve exits skull through sphenorbital fissure (0); present (1) (Novacek, 1986; Thewissen and Domning, 1992).

• X [37] 031. Mastoid foramen.—-Present, skull in posterior view (0); absent (1) (de?nition of structure follows MacPhee, 1994).

• X [38] 032. Post-temporal canal (for arteria diploetica magna, also called pericranial foramen).—-Present, occurs at petrosal/squamosal suture with skull in posterior view, the canal continuing within the pet- rosal/squamosal suture (0); absent (1) (Wible, 1990; MacPhee, 1994).

• X [39] 033. Preglenoid process.—-Absent(0);present, forms trans- verse, ventrally projecting ridge at anterior edge of glenoid fossa (1) (modi?ed from Thewissen, 1994). Cranial Characters

• X [40] 034. Supraorbital process.—-Absent,region over orbit does not project laterally from sagittal plane (0); present, laterally elongate and tabular (1) (derived from Barnes, 1984).

• X [41] 035. Postorbital bar (ordered).—-Absent (0); present and almost complete (1);

• X [42] …. present and complete (2).

• X [43] 036. Contact of frontal and maxilla in orbit.—-Absent (0); present (1).

• X [44] 037. Lacrimal tubercle.—-Absent (0); present, situated on anterior edge of orbit adjacent to the lacrimal fora- men (1) (Novacek, 1986).

• X [45] 038. Palatine ?ssure.—-Small (0); large (1).

• X [46] 039. Posterior margin of external nares (ordered).—- Anteriorto or overthe canines, or at anterior edge of premaxilla if canine not present (0); between P1 and P2 (1);

• X [47] …. posterior to P2 (2) (Geisler and Luo, 1998).

• X [48] 040. Mandibular foramen.—-Small, maximum height of opening #25% the height of the mandible at M3 (0); large, continuous with a large posterior fossa, maximum height $50% the height of the mandible at M3 (1) (modified from Thewissen, 1994; Geisler and Luo, 1998).

• T [49] 041. Height of mandibular condyle relative to dentition (ordered).—-Below level of the dentition (0); even with superior aspect of dentition (1);

• T [50] ….substantially superior to dentition (2).

• X [51] 042. Angle of dentary.—-Distal end at same level as ventral edge of dentary below molars (0); forms distinct flange that projects posteroventrally well below ventral edge of dentary (1) (modi?ed from Gentry and Hooker, 1988).

• N [52] 043. Upper incisors.—-Present (0); absent (1).

• T [53] 044. Premaxillae.—-Short with incisors arranged in trans- verse arc (0); elongate, incisors aligned longitu- dinally with intervening diastemata (1) (modi?ed from Prothero et al., 1988; Thewissen, 1994).

• N [54] 045. Embrasure pits on palate.—-Absent (0); present (1) (modi?ed from Thewissen, 1994).

• N [55] 046. P1 (ordered).—-Absent (0); present, one-rooted (1);

• T [56] …. present, two-rooted (2) (Zhou et al., 1995).

• T [57] 047. P3 roots.—-Three (0); two (1) (Zhou et al., 1995).

• T [58] 048. P4 protocone.—-Present (0); absent (1) (Thewissen, 1994).

• T [59] 049. P4 paracone.—-Equal or subequal to height of para- cone of M1 (0); greater than twice the height of M1 paracone (1) (Thewissen, 1994).

• T [60] 050. P4 metacone.—-Absent (0); present (1) (Thewissen, 1994).

• I [61] 051. Stylar shelf.—-Present (0); absent (1).

• T [62] 052. M2 metacone (ordered).—-Distinct cusp, sub-equal to paracone (0); distinct cusp approximately half the size of the paracone (1);

• N [63] …. highly reduced, indistinct from paracone (2) (Zhou et al., 1995).

• N [64] 053. M1 parastyle (ordered).—- Absent (0); weak (1);

• N [65] … mod- erate to strong (2) (O’Leary, 1998a).

• I [66] 054. Lingual cingulum on M2 .—-Present (0); absent (1) (O’Leary, 1998a).

• I [67] 055. Ectocingula.—-Present (0); absent (1) (O’Leary, 1998a).

• N [68] 056. M1 -M2 hypocone.—-Absent (0); present (1) (Thewis- sen and Domning, 1992; O’Leary, 1998a).

• T [69] 057. Trigon basin (ordered).—-Broad (0); somewhat nar- row (1);

• T [70] …. very narrow (2) (modified from Thewissen, 1994; O’Leary, 1998).

• N [71] 058. Paraconule on M2 .—-Absent (0); present (1) (O’Leary, 1998a).

• N [72] 059. Metaconule on M2 .—-Absent(0); present(1) (O’Leary, 1998a).

• I [73] 060. M3 size (ordered).—-Present, larger than M2 (0); present, approximately equal to M2 (1);

• N [74] …. present, small, maximum mesiodistal length than < 60% the length of M2 (2);

• N [75] …. absent (3) (modi?ed from Zhou et al., 1995).

• X [76] 061. Lower canines.—-Approximately twice as large as in- cisors or larger (0); subequal in size to incisors, well separated from postcanine teeth (1).

• X [77] 062. Lower incisors.—-Small and vertical (0); tusk-like (1).

• T [78] 063. Lower canine cross-section shape.—-Oval (0); triangular (1) (Gentry and Hooker, 1988).

• N [79] 064. P1.—-Present (0); absent (1) (Zhou et al., 1995). [Assume that loss is secondary]

• N [80] 065. P3 metaconid.—-Absent (0); present (1) (Thewissen and Domning, 1992).

• N [81] 066. P4 metaconid.—-Absent (0); present (1) (Thewissen and Domning, 1992).

• N [82] 067. Lingual cingulid on lower molars.—-Poorly de?ned or absent (0); continuous from mesial to distal extreme (1) (O’Leary, 1998a).

• N [83] 068. M1 paraconid.—-Present (0); absent (1) (derived from O’Leary, 1998a).

• T [84] 069. M1 paraconid/paracristid position.—-Cusp lingual or paracristid winding lingually (or both) (0); cusp anterior or paracristid (or both) straight mesiodistally, sometimes poorlydeveloped (1) (derivedfrom O’Leary, 1998a).

• N [85] 070. M2 paraconid.—-Present (0); absent (1) (derived from O’Leary, 1998a).

• T [86] 071. M2 paraconid/paracristid position.—-Cusp lingual or paracristid (or both) winding lingually (0); cusp anterior or paracristid (or both) straight mesiodistally, sometimes poorly developed (1) (O’Leary, 1998a).

• N [87] 072. M3 paraconid.—-Present (0); absent (1) (O’Leary, 1998a).

• T [88] 073. M3 paraconid/paracristid position.—-Cusp lingual or paracristid winding lingually (or both) (0); cusp anterior or paracristid (or both) straight mesiodistally, sometimes poorly developed (1) (O’Leary, 1998a).

• N [89] 074. M1 metaconid.—-Present, forms distinct cusp (0); absent or occasionally present as swelling on lingual side of protoconid (1) (Thewissen, 1994; Zhou et al., 1995).

• N [90] 075. M2 metaconid.—-Present, forms distinct cusp (0); absent or occasionally present as swelling on lingual side of protoconid (1) (modi?ed from Zhou et al., 1995).

• N [91] 076. M3 metaconid.—-Present, forms distinct cusp (0); absent or occasionally present as swelling on lingual side of protoconid (1) (modi?ed from Zhou et al., 1995).

• N [92] 077. M1 -M2 hypoconulid.—-Absent (0); present (1) (The- wisen, 1994).

• T [93] 078. Molar protoconid.—-Subequal to height of talonid (0); closer to twice height of talonid or greater (1) (O’Leary, 1998a).

• X [94] 079. Protoloph.—-Absent (0); present (1).

• X [95] 080. Metaloph.—-Absent (0); present (1).

• X [96] 081. Metalophid formation on anterior aspect of lower teeth (between protoconid and metaconid).—-Absent (0); present (1).

• X [97] 082. Hypolophid formation on posterior aspect of lower teeth (between entoconid and hypoconid).—-Absent (0); present (1) (Thewissen and Domning, 1992).

• X [98] 083. Reentrant grooves.—-Proximal (0); absent (1); distal (2) (Thewissen, 1994; O’Leary, 1998a).

• X [99] 084. Molar talonid.—-Basined, or slightly basined, hypoconid and entoconid present (0); reduced (1) (modi?ed from Thewissen, 1994).

• X [100] 085. M3 hypoconulid.—-Long, protrudes as separate distal lobe (0); short, does not protrude substantially beyond rest of talonid (1); absent (2) (Thewissen, 1994).

• X [101] 086. dp4 .—-Resembles M1 (0); six-cusped, additional neomorphic cusp on paracristid (1) (Gentry and Hooker, 1988; Luckett and Hong, 1998).

• X [102] 087. Elongate shearing facets on lower molars extending below the gum line.—-Absent (0); present (1). Postcranial Characters

• X [103] 088. Cervical vertebrae.—-Long, length of centrum greater than or equal to the centra of the anterior thoracics (0); short, length shorter than centra of anterior tho- racics (1); very long (2) (derived from Gingerich et al., 1995).

• X [104] 089. Articulation between sacral vertebrae and ilium of pelvis (ordered).—-Broad area of articulation between pelvis and S1 and sometimes S2 (0); narrow articulation of pelvis with end of transverse process of S1 (1);

• X [105] …. articulation absent (2) (Geisler and Luo, 1998).

• X [106] 090. Number of sacral vertebrae (ordered).—-One (0); two or three (1);

• X [107] …. four (2);

• X [108] …. five or six (3); cannot be scored for taxa that lack an articulation between the verte- bral column and the ilium (Thewissen and Domning, 1992; Gingerich et al., 1995).

• X [109] 091. Scapular spine.—-Bears large acromion pro- cess that overhangs glenoid fossa (0); scapular spine with acromion process small or absent, does not encroach upon glenoid fossa (1); acromion process large, directed anteriorly and does not encroach upon the glenoid fossa (2) (O’Leary and Rose, 1995b; Geisler and Luo, 1998).

• X [110] 092. Entepicondyle of humerus.—-Wide, $50% width of ul- nar and radial articular facets (0); narrow, #25% width of ulnar and radial articular facets (1) (de- rived from O’Leary and Rose, 1995b; Geisler and Luo, 1998).

• X [111] 093. Entepicondylar foramen.—-Present (0); absent (1) (Thewissen and Domning, 1992).

• X [112] 094. Length of olecranon process.—-Short, < 10% total ul- nar length (0); long, > 20% total ulnar length (1) (derived from O’Leary and Rose, 1995b).

• X [113] 095. Olecranon fossa.—-Shallow (0); deep, sometimes perforate (1) (modi?ed from O’Leary and Rose, 1995b).

• X [114] 096. Proximalend of radius.—-Single fossa fortrochlea and capitulum of humerus (0); two fossae, one forthe ca- pitulum and one for the medial edge of trochlea (1); three fossae, same as (1) but with additional fossa for the lateral lip of the humeral articular surface (2) (O’Leary and Rose, 1995b; Geisler and Luo, 1998).

• X [115] 097. Distal articular surface of radius.—-Single concave fossa (0); split into scaphoid and lunate fossae (1); convex to flat, distinct facets (2) (derived from O’Leary and Rose, 1995b).

• X [116] 098. Centrale.—-Present (0); (1) absent (Thewissen, 1994).

• X [117] 099. Manus.—-Mesaxonic, axis of symmetry passes along center of digit 3 (0); paraxonic, axis of symmetry passes between digits three and four (1).

• X [118] 100. Height of greater trochanter of femur (ordered).—- Inferiorto head of femur(0); approximately equal to head of femur (1);

• X [119] …. above head of femur (2) (derived from O’Leary and Rose, 1995b).

• N [120] 101. Third trochanter of femur (ordered).—-Present (0); highly reduced (1);

• N [121] …. absent(2) (derived from O’Leary and Rose, 1995b).

• T [122] 102. Patellar articular surface on femur.—-Wide (0); narrow (1).

• I [123] 103. Astragalar canal.—-Present (0); absent (1) (derived from Shoshani, 1986).

• T [124] 104. Navicular facet of astragalus (ordered).—- Convex (0); saddle-shaped(1);

• T [125] …. trochleated (2)(modi?ed from Thewissen and Domning, 1992; O’Leary and Rose, 1995b).

• T [126] 105. Sustentacular width.—-Narrow, #50% width of the astragalus (0); wide, $70% width of the astragalus (1) (Geisler and Luo, 1998).

• N [127] 106. Contact of distal astragalus with cuboid (ordered).—-Absent (0); present but small, contact # 30% of the width of the distal articular surface (1);

• T [128] …. present and large, contact $40% of the articular surface (2) (O’Leary and Rose, 1995b; Geisler and Luo, 1998).

• I [129] 107. Lateral process of astragalus.—-Present (0); absent (1) (Schaeffer, 1956).

• T [130] 108. Pes(ordered).—-Mesaxonic, axis of symmetry of foot passes along center of digitthree (0); paraxonic, axis lies between digits three and four (1);

• T [131] …. mesaxonic, axis passes along center of digit four (2) (derived from Gingerich et al., 1990; Thewissen, 1994; our interpretation of this character differs from that of these previous authors).

• T [132] 109. Proximal astragalus (ordered).—-Nearly flat to slightly concave (0); grooved, depth of trochlea < 25% width of trochlea (1);

• T [133] …. deeply grooved, depth of trochlea > 30% width of trochlea (2) (derived from Schaeffer, 1947; O’Leary and Rose, 1995b).

• N [134] 110. Cuboid and navicular.—-Unfused (0); fused (1) (Webb and Taylor, 1980).

• X [135] 111. Proximal fusion of third and fourth metatarsals.—- Absent (0); present (1).

• X [136] 112. First metatarsal (ordered).—-Unreduced, length > 50% length of third metatarsal (0); reduced, length < 50% length of third metatarsal(1);

• X [137] …. highly reduced in form of nodule or small splint or absent (2).

• X [138] 113. Second metatarsal (ordered).—-Unreduced, length $50% length of third metatarsal(0); reduced, length #50% length of third metatarsal (1);

• X [139] …. highly reduced in form of nodule, small splint or completely absent (2).

• X [140] 114. Fifth metatarsal (ordered).—-Unreduced, length $50% length of third metatarsal (0); reduced, length #50% length of third metatarsal (1);

• X [141] …. highly reduced in form of nodule or small splint or absent (2).

• X [142] 115. Ventral border of distal phalanges.—-Curved inferiorly (0); straight (1) (derived from MacLeod and Rose, 1993).

• X [143] 116. Distal phalanges in dorsal view.—-Phalanx com- pressed transversely (0); broad transversely, each phalanx bilaterally symmetricalwith central antero- posterior axis (1); broad transversely, each phalanx asymmetrical (2) (derived from MacLeod and Rose, 1993). Soft Tissue Characters

• X [144] 117. Stomach epithelium.—-Nonglandular(strati?edsqua- mous epithelium) (0); composite (nonglandular and glandular mucosa) (1); discoglandular (small patches of glandular epithelium surrounded by nonglandular mucosa) (2); glandular (mucosa with glands) (3) (derived from Langer, 1988).

• X [145] 118. Lumen.—-Unilocular (stomach with simple cham- ber only) (0); plurilocular (lumen of stomach sub- divided by folds into two or more chambers) (1) (derived from Langer, 1988).

• X [146] 119. Diverticulation of stomach lumen (blind tubes or sacs branching from the main gastric lumen).—-Absent (0); present (1) (derived from Langer, 1988).

• X [147] 120. Cavernous tissue of penis.—-Abundant (0); sparse (1) (derived from Slijper, 1936; Thewissen, 1994).

• X [148] 121. Hair.—-Abundant to common on body (0); almost completely absent (1) (Gatesy, 1997).

• X [149] 122. Sebaceous glands.—-Present (0); absent (1) (Gatesy, 1997).

• X [150] 123. Primary bronchi of lung.—-Two (0); three (1) (Thewis- sen, 1994)

63 taxa, 317 characters, of which 9 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 87, 89, 91, 92, 93, 94, 99, 100, 118

• Neomorphic:

Details:

• T [118] Ancestral state considered ambiguous, so treated as transformational

58 taxa, 317 characters, of which 9 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 87, 89, 91, 92, 93, 94, 99, 100, 118

• Neomorphic:

Details:

• T [118] Ancestral state considered ambiguous, so treated as transformational

34 taxa, 107 characters, of which 55 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 20, 23, 24, 25, 31, 33, 40, 56, 57, 65, 69, 70, 72, 74, 76, 78, 80, 81, 86, 89, 90, 99, 103, 106

• Neomorphic: 15, 16, 17, 18, 19, 21, 22, 36, 58, 59, 60, 61, 62, 71, 73, 75, 77, 79, 82, 84, 104

Details:

• X [1] Prostomium: distinct 0; fused to peristomium 1; frontal edge fused to peristomium 2.

• X [2] Shape of prostomium: lobed-ridge-shaped 0; hood-like flattened 1; shovel-shaped 2; limited to palps 3; lobes 4.

• X [3] Peristomium: lips only 0; ring or rings 1; extended upper and lower lips 2.

• T [4] Peristomial rings: one ring 0; two rings 1.

• T [5] Margin of anterior peristomial ring: low external 0; ventral as triangular lobe 1.

• T [6] Peristomial ring: smooth 0; with collar 1.

• X [7] Palps: absent 0; present 1.

• T [8] Appearance of palps: sensory ventral 0; grooved 1.

• T [9] Origin of palps: prostomial 0; peristomial 1.

• T [10] Adult location of palps: outside mouth 0; inside mouth 1.

• T [11] Contractibility of palps: minimal 0; extensive 1.

• T [12] Prostomial palps: paired 0; multiple 1; crown 2.

• T [13] Radiolar lobes: separated 0; dorsally fused 1.

• T [14] Radiole fusion: separate 0; with palmate membrane 1.

• N [15] Radiolar flanges: absent 0; present 1.

• N [16] Radiolar skeleton: absent 0; present 1.

• N [17] Parallel lamellae: absent 0; present 1.

• N [18] Peristomial palps: paired 0; single 1; multiple 2.

• N [19] Peristomial palp pinnules: absent 0; present 1.

• T [20] Peristomial palp pinnules: multicelled 0; single-celled 1.

• N [21] Fused multiple peristomial palps: absent 0; present 1.

• N [22] Obturaculum: absent 0; present 1.

• T [23] Obturaculum size: short 0; long 1.

• T [24] Muscular arrangement in obturaculum: parasagittal 0; frontal 1.

• T [25] Dorsal groove in the obturaculum: grooved 0; ridged 1.

• X [26] Retractable head: absent 0; present 1.

• X [27] Structure of first segment: dorsally limited 0; fused to head 1; elongate 2; similar to following 3.

• X [28] Segment 2 forming elongate trunk: absent 0; present 1.

• X [29] Metameric papillae on segment 2: absent 0; present 1.

• X [30] Girdle on segment 2: absent 0; present 1.

• T [31] Uncinal girdle on segment 2: median 0; posterior 1.

• X [32] Papillae on posterior part of segment 2: absent 0; present 1.

• T [33] Arrangement of papillae on posterior part of segment 2: line or rows 0; scattered 1.

• X [34] Vestimentum: absent 0; present 1.

• X [35] Frenulum: absent 0; present as continuous ridge 1; scattered plaques only

• N [36] Diaphragm with distinct external groove: absent 0; present 1.

• X [37] Lateral lobes on anterior segments: absent 0; present 1.

• X [38] Thoracic membrane on anterior segments: absent 0; present 1.

• X [39] Ventral glandular area on anterior segments: absent 0; present 1.

• T [40] Shape of ventral glandular area on anterior segments: ventral central pads 0; ventral annullae 1.

• X [41] Epidermal papillae: ventral central pads 0; ventral anullae 1.

• X [42] Building organ: absent 0; present 1.

• X [43] Notochaetae on segment 1: absent 0; present 1.

• X [44] Notochaetae on segment 2: absent 0; present 1.

• X [45] Notochaetae on segment 3: absent 0; present 1.

• X [46] Notochaetae on segment 4: absent 0; present 1.

• X [47] Neurochaetae on segment 1: absent 0; present 1.

• X [48] Neurochaetae on segment 2: absent 0; present 1.

• X [49] Neurochaetae on segment 3: absent 0; present 1.

• X [50] Neurochaetae on segment 4: absent 0; present 1.

• X [51] Neurochaetae on segment 5: absent 0; present 1.

• X [52] Neurochaetae on segment 6: absent 0; present 1.

• X [53] Neurochaetae on segment 7: absent 0; present 1.

• X [54] Neurochaetae on segment 8: absent 0; present 1.

• X [55] Branchiae: absent 0; present 1.

• T [56] Shape of branchiae: simple filaments 0; branching numerous lamellae 1.

• T [57] Arrangement of branchiae: segmentally 0; grouped on single segment 1.

• N [58] Branchiae on segment 1: absent 0; present 1.

• N [59] Branchiae on segment 2: absent 0; present 1.

• N [60] Branchiae on segment 3: absent 0; present 1.

• N [61] Branchiae on segment 4: absent 0; present 1.

• N [62] Branchiae on segment 5: absent 0; present 1.

• X [63] Chaetal inversion: absent 0; present 1.

• X [64] Parapodia: projecting neuropodia 0; similar rami 1; noto- and neuropodia differing; none 2.

• T [65] Anterior neuropodia segment 2–8: short, truncate cylinders 0; low ridges or pads 1; tori 2; with lobes or lamellae 3; chaetae from body wall 4.

• X [66] Posterior neuropodia: short, truncate cylinders 0; low ridges or pads 1; tori 2; with lobes or lamellae 3; straight from body wall 4.

• X [67] Dorsal half of posterior body segments: with parapodia and chaetae 0; smooth 1.

• X [68] Hooks/ uncini: absent 0; present 1.

• T [69] Arrangement of hooks/ uncini: single row 0; two rows 1; dense fields 2.

• T [70] Hook projection: projecting beyond tip of parapodia 0; low 1.

• N [71] Capitium: absent 0; present 1.

• T [72] Organization of capitium: single column 0; in two or more columns 1; arched rows over rostrum 2; bidentate 3.

• N [73] Rostrum: absent 0; present 1.

• T [74] Size of rostrum: small tooth 0; large fang 1.

• N [75] Subrostrum (breast of hooks/ uncini): absent 0; present 1.

• T [76] Size of breast of subrostrum: extending less than capitium or rostrum 0; extending further than capitium or rostrum 1.

• N [77] Subrostral process of hooks/ uncini: absent 0; present 1.

• T [78] Subrostral process of hooks/ uncini. appendix for muscle attachment 0; teeth in same direction as capitium teeth 1; teeth oppose capitium teeth 2.

• N [79] Manubrium of hooks/ uncini: absent 0; present 1.

• T [80] Length manubrium of hooks/ uncini: shorter than remainder of hook / uncinus 0; longer than remainder of hook / uncinus 1.

• T [81] Uncinal hook forms: hooks only 0; found on some segments 1.

• N [82] Hoods of hooks/ uncini protection: absent 0; present 1.

• X [83] Chaetae in posterior segments: mixed capillaries or spines 0; hooks/ uncini only in notopodial position 1; hooks/ uncini in noto- and neuropodial positions 2; hooks/ uncini only in neuropodial position 3.

• N [84] Companion chaetae: absent 0; present 1.

• X [85] Notopodial spines: absent 0; present 1.

• T [86] Notopodial spines: along major part of body 0; projecting forward as paleae 1; in few anterior segments only 2.

• X [87] Caudal region: absent 0; present 1.

• X [88] Opisthosoma: absent 0; present 1.

• T [89] Length opisthosoma: short 0; long 1.

• T [90] Opisthosomal chaetae: parapodial 0; single long rows 1.

• X [91] Inverted faecal groove: absent 0; present 1.

• X [92] Gular membrane: absent 0; present 1.

• X [93] Subdermal intraepidermal nerve cord: absent 0; present 1.

• X [94] Lateral organs: absent 0; present 1.

• X [95] Mouth: axial proboscis 0; simple tube 1; ventral buccal organ 2.

• X [96] Gut: straight 0; looped 1.

• X [97] Occluded gut: absent 0; present 1.

• X [98] Anterior segmental organs: not differentiated from rest 0; very enlarged pair 1; absent 2; excretory series 3.

• T [99] Single anterior pair of nephridia: two openings 0; single opening 1.

• X [100] Genital papillae: none apparent 0; on chaetiger 5–6 1.

• X [101] Heart bodies: absent 0; present 1.

• X [102] Tube: absent 0; present 1.

• T [103] Tube construction: sand and or mucus 0; chitin protein complex 1; calcareous 2.

• N [104] Tube attachment to substrate: absent 0; present 1.

• X [105] Sperm packaging: absent 0; present 1.

• T [106] Sperm packaging: spermatophores 0; spermatozeugmata 1.

• X [107] Larvae: planktotrophic trochophore 0; lecithotrophic trochophore 1; mitraria 2.

36 taxa, 52 characters, of which 13 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 8, 13, 16, 19, 29, 33, 35, 37, 40, 41, 42, 51

• Neomorphic: 43

Details:

• X [1] Apterous exule. Body shape: (0) oblong; (1) round.

• X [2] Fundatrix or apterous exule. Relative length of antennae to body length: (0) < 0.2; (1) > 0.2.

• X [3] Emigrant, sexupara or alate exule. Relative length of antennae to body length: (0) < 0.4; (1) > 0.4.

• X [4] Fundatrix. Number of antennal segments: (0) three; (1) four; (2) five; (3) six.

• X [5] Apterous exule. Number of antennal segments: (0) three; (1) four; (2) five; (3) six.

• X [6] Emigrant, sexupara or alate exule. Number of antennal segments: (0) five; (1) six.

• X [7] Emigrant, sexupara or alate exule. Relative length of antennal segment III to segments IV, V and VI combined (for six-segmented antennae): (0) < 0.8; (1) 0.8–1.2; (2) > 1.2.

• T [8] Emigrant, sexupara or alate exule. Relative length of antennal segment IV to segment V (for six-segmented antennae): (0) < 0.7; (1) > 0.7.

• X [9] Emigrant, sexupara or alate exule. Development of processus terminalis: (0) undeveloped (the length is shorter than or almost as long as the width) (Fig. 3A, B); (1) developed (the length is much larger than the width).

• X [10] Emigrant, sexupara or alate exule. Shape of secondary rhinarium: (0) annular (secondary rhinaria usually cover much more than half the circumference from the ventral side); (1) transverse oval (secondary rhinaria cover less than or nearly half the circumference); (2) round (secondary rhinaria cover much less than half the circumference).

• X [11] Emigrant, sexupara or alate exule. Colour of secondary rhinarium: (0) bright, or paler than background; (1) pigmented.

• X [12] Emigrant, sexupara or alate exule. Margins of secondary rhinarium: (0) not rimmed; (1) rimmed.

• T [13] Emigrant, sexupara or alate exule. Projection of secondary rhinarium: (0) hardly projected, interhinarial parts not constricted (Fig. 3C, D); (1) projected, interhinarial parts constricted.

• X [14] Emigrant, sexupara or alate exule. Secondary rhinarium on antennal segment V : (0) absent; (1) present.

• X [15] Emigrant, sexupara or alate exule. Secondary rhinarium on antennal segment VI: (0) absent (Fig. 3A, E); (1) present.

• T [16] Emigrant, sexupara or alate exule. Number and development of secondary rhinaria on antennal segment VI (if present): (0) one to three, rudimentary (secondary rhinaria are much fewer and smaller than those on other segments); (1) one to three, developed (secondary rhinaria are much fewer than those on other segments, but are almost the same in size); (2) more than three, developed (secondary rhinaria are almost the same as those on other segments in number and size).

• X [17] Emigrant, sexupara or alate exule. Relative length of ultimate rostral segment to second segment of hind tarsus: (0) < 0.8; (1) 0.8–1.2; (2) >1.2.

• X [18] Fundatrix or apterous exule. Membranous slit between first and second tarsal segments: (0) absent (first and second tarsal segments completely fused); (1) present partially or entirely (Fig. 4D, E).

• T [19] Fundatrix or apterous exule. Development of membranous slit on tarsi (if present): (0) incomplete; (1) complete.

• X [20] Apterous exule. Stout seta with a large prominent socket on the ventral side of fore tarsus: (0) absent (Fig. 4A, D, E); (1) present, weakly developed (the socket situated at a ventro-middle point, not reaching the base of claws); (2) present, well developed (the socket situated at a ventro-anterior point, reaching or exceeding the base of claws).

• X [21] Emigrant, sexupara or alate exule. Number of ventroanterior setae on the first segment of fore tarsi: (0) two; (1) three; (2) four; (3) more than four.

• X [22] Emigrant, sexupara or alate exule. Number of ventroanterior setae on the first segment of middle tarsi: (0) two; (1) three; (2) four; (3) more than four.

• X [23] Emigrant, sexupara or alate exule. Number of ventroanterior setae on the first segment of hind tarsi: (0) two; (1) three; (2) four; (3) more than four.

• X [24] Apterous exule. Number of ventro-anterior setae on the first segment of fore tarsi: (0) two; (1) three; (2) more than three.

• X [25] Apterous exule. Number of ventro-anterior setae on the first segment of middle tarsi: (0) two; (1) three; (2) more than three.

• X [26] Apterous exule. Number of ventro-anterior setae on the first segment of hind tarsi: (0) two; (1) three; (2) more than three.

• X [27] Emigrant, sexupara or alate exule. Media of fore wings: (0) unbranched; (1) once branched.

• X [28] Emigrant, sexupara or alate exule. Oblique veins of hind wings: (0) one vein, rarely with an additional indistinct vein; (1) two distinct veins.

• T [29] Emigrant, sexupara or alate exule. The bases of two oblique veins (if two oblique veins are present): (0) distinctly separated; (1) fused or very close to each other. Abdominal sclerite

• X [30] Emigrant. Abdominal sclerites on anterior tergites: (0) absent; (1) present.

• X [31] Sexupara or alate exule. Abdominal sclerites on anterior tergites: (0) absent; (1) present.

• X [32] Emigrant. Siphunculi: (0) absent; (1) present.

• T [33] Emigrant. Shape of siphunculi (if present): (0) small pores, not surrounded by setae nor sclerites; (1) small pores, surrounded by a few setae and indistinct sclerites; (2) large pores, surrounded by several setae and distinct sclerites.

• X [34] Apterous or alate exule, or sexupara. Siphunculi: (0) absent; (1) present.

• T [35] Apterous or alate exule, or sexupara. Shape of siphunculi (if present): (0) small pores, not surrounded by setae nor sclerites; (1) small pores, surrounded by a few setae and indistinct sclerites; (2) large pores, surrounded by several setae and distinct sclerites.

• X [36] Fundatrix. WGPs: (0) absent; (1) present, either indistinct or distinct.

• T [37] Fundatrix. Development of WGPs (if present): (0) indistinct, without clear borders; (1) distinct, with clear borders.

• X [38] Emigrant. WGPs on abdomen: (0) absent; (1) present.

• X [39] Sexupara or alate exule. WGPs on abdomen: (0) absent; (1) present.

• T [40] All morphs with WGPs. Size of cells in a WGP: (0) uniform (WGPs consist of cells of almost the same size); (1) not uniform (WGPs consist of cells of different size).

• T [41] All morphs with WGPs. Central fields of a WGP: (0) absent, or indistinct; (1) present and distinct.

• T [42] All morphs with WGPs. Number and shape of central fields of a WGP (if present): (0) several, circular; (1) single, circular; (2) single, elongate, sometimes fragmented into several small ones.

• N [43] All morphs with WGPs. Setae on a WGP: (0) absent; (1) present.

• X [44] Apterous exule. Arrangement of WGPs on prothorax: (0) absent; (1) present on the lateral position; (2) present on the spinal and lateral positions.

• X [45] Apterous exule. Arrangement of WGPs on meso- and metathorax: (0) absent; (1) present on the lateral position; (2) present on the spinal and lateral positions; (3) present on the pleural and lateral positions; (4) present on the spinal, pleural and lateral positions.

• X [46] Apterous exule. Arrangement of WGPs on abdominal segments I and II: (0) absent; (1) present on the lateral position; (2) present on the spinal and lateral positions; (3) present on the spinal, pleural and lateral positions.

• X [47] Apterous exule. Arrangement of WGPs on abdominal segments III and IV : (0) absent; (1) present on the lateral position; (2) present on the spinal and pleural positions; (3) present on the spinal and lateral positions; (4) present on the spinal, pleural and lateral positions.

• X [48] Apterous exule. Arrangement of WGPs on abdominal segment V : (0) absent; (1) present on the lateral position; (2) present on the spinal and pleural positions; (3) present on the spinal and lateral positions; (4) present on the spinal, pleural and lateral positions.

• X [49] Apterous exule. Arrangement of WGPs on abdominal segment VI: (0) absent; (1) present on the lateral position; (2) present on the spinal and pleural positions; (3) present on the spinal and lateral positions; (4) present on the pleural and lateral positions; (5) present on the spinal, pleural and lateral positions.

• X [50] Apterous exule. Arrangement of WGPs on abdominal segment VII: (0) absent; (1) present on the spinal position; (2) present on the lateral position; (3) present on the spinal and pleural positions; (4) present on the spinal and lateral positions; (5) present on the pleural and lateral positions; (6) present on the spinal, pleural and lateral positions.

• T [51] Apterous exule. Fusion of WGPs on abdominal segment VII (if spinal and pleural WGPs, or pleural and lateral WGPs are present): (0) not fused; (1) spinal and pleural pairs fused; (2) pleural and lateral pairs fused.

• X [52] Apterous exule. Arrangement of WGPs on abdominal segment VIII: (0) absent; (1) one spinal pair; (2) one at the central position; (3) one expanding widely on the whole tergite

23 taxa, 109 characters, of which 34 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 10, 14, 16, 17, 21, 25, 30, 46, 59, 76, 81, 82, 86, 89, 100, 101, 102, 104, 105, 106, 108, 109

• Neomorphic: 2, 5, 18, 22, 26, 39, 63, 66, 84, 91, 103, 107

52 taxa, 58 characters, of which 29 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 10, 11, 13, 14, 16, 18, 22, 23, 24, 25, 28, 29, 32, 37, 38, 44, 50, 57, 58

• Neomorphic: 19, 27, 30, 31, 33, 34, 35, 36, 46, 48

Details:

• X [1] Peripheral tentacles: 0 - absent, 1 - present

• X [2] Nuchal tentacles: 0 - absent, 1 - present

• X [3] Branched tentacles: 0 - absent, 1 - present

• X [4] Tentacles along introvert: 0 - absent, 1 - present

• X [5] Nephridia: 0 - paired, 1 - single

• X [6] Nephridial attachment: 0 - mostly unattached, 1 - at least 50% attached

• X [7] Nephridial shape: 0 - unilobed, 1 - bilobed

• X [8] Position of nephridiopores: 0 - anterior to anus, 1 - same level as anus, 2 - posterior to anus

• X [9] Coelomic extensions in body wall: 0 - absent, 1 - present

• T [10] Type of coelomic extensions: 0 - sacs, 1 - canals

• T [11] Orientation of coelomic canals: 0 - longitudinal, 1 - in bands

• X [12] Longitudinal musculature: 0 - distinct, 1 - anastomosing

• T [13] Number of longitudinal muscle bands: 0 - commonly < 25; 1 - 25-40, 2 - > 40

• T [14] Longitudinal muscle bands: 0 - distinct, 1 - anastomosing

• X [15] Circular musculature: 0 - continuous sheet, 1 - in bands

• T [16] Circular muscle bands: 0 - distinct, 1 - anastomosing

• X [17] Dorsal shield: 0 - absent, 1 - present

• T [18] Shape of dorsal shield: 0 - flat, 1 - cone-shaped

• N [19] Grooves in dorsal shield: 0 - absent, 1 - present

• X [20] Pineapple shield: 0 - absent, 1 - present

• X [21] Spindle muscle: 0 - absent, 1 - present

• T [22] Anterior attachment of spindle muscle: 0 - body wall, 1 - on rectum

• T [23] Level of attachment on body wall: 0 - anterior to anus, 1 - same level as anus, 2 - posterior to anus

• T [24] Anterior roots of spindle muscle: 0 - one, 1 - two or more

• T [25] Posterior attachment of spindle muscle: 0 - posterior body wall, 1 - within gut coil

• X [26] Hooks on introvert in adults: 0 - absent, 1 - present

• N [27] Hooks in rings: 0 - absent, 1 - present

• T [28] Number of rings of hooks: 0 - < 50, 1 - > 50

• T [29] Hook arrangement: 0 - some scattered, 1 - entirely in rows

• N [30] Bidentate hooks: 0 - absent, 1 - present

• N [31] Secondary tooth on hook: 0 - absent, 1 - present

• T [32] Shape of secondary tooth: 0 - blunt, 1 - pointed

• N [33] Hook interior: Anterior clear triangle: 0 - absent, 1 - present

• N [34] Hook interior: Clear streak: 0 - absent, 1 - present

• N [35] Hook interior: Crescent-shaped space: 0 - absent, 1 - present

• N [36] Posterior basal structures: 0 - absent, 1 - present

• T [37] Posterior basal structure type: 0 - warts, 1 - rootlets

• T [38] Angle of hook: 0 - < 90%, 1 - > 90%

• X [39] Pyramidal hooks: 0 - absent, 1 - present

• X [40] Conical hooks: 0 - absent, 1 - present

• X [41] Location of anus: 0 - on anterior trunk, 1 - on introvert

• X [42] Pigmented introvert bands: 0 - absent, 1 - present

• X [43] Contractile vessel villi: 0 - absent, 1 - 1 present

• T [44] Type of contractile vessel villi: 0 - digitiform, 1 - elongate tubules

• X [45] Introvert retractor muscles in adult: 0 - two pairs, 1 - less than two pairs

• N [46] Fusion of dorsal retractors: 0 - not fused (< 10%), 1 - partially fused, 2 - completely fused

• X [47] Fusion in ventral retractors: 0 - not fused (< 10%), 1 - partially fused, 2 - completely fused

• N [48] Fusion of dorsal and ventral retractors: 0 - absent, 1 - present

• X [49] Retractor column: 0 - absent, 1 - present

• T [50] Origin of dorsal retractor muscles: 0 - anterior 1/3 of body, 1 - middle 1/3 of body, 2 - posterior 1/3 of body

• X [51] Origin of ventral retractor muscles: 0 - anterior 1/3 of body, 1 - middle 1/3 of body; 2 - posterior 1/3 of body

• X [52] Protractor muscle: 0 - absent, 1 - present

• X [53] Ratio of introvert/trunk length: 0 - < 0.75, 1 - 0.75-2, 2 - > 2

• X [54] Holdfast papillae with hardened borders: 0 - absent, 1 - present

• X [55] Caudal appendage: 0 - absent, 1 - present

• X [56] Nuchal organ: 0 - pit, 1 - ciliated band, 2 - patch or patches

• T [57] Number of nuchal patches: 0 - one, 1 - two

• T [58] Shape of nuchal patch: 0 - triangular, 1 - bilobed, 2 - multi-lobed

59 taxa, 202 characters, of which 33 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 15, 25, 36, 44, 78, 84, 113, 118, 137, 140, 144, 165, 200, 201

• Neomorphic: 6, 31, 45, 66, 73, 74, 85, 99, 107, 114, 119, 126, 141, 145, 153, 164, 183, 192, 202

Details:

• T [200] Attachment, not presence/absence

• T [201] Attachment, not presence/absence

23 taxa, 57 characters, of which 25 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 10, 39, 42, 45, 51, 56

• Neomorphic: 13, 14, 15, 28, 29, 30, 33, 40, 41, 43, 46, 47, 48, 49, 50, 52, 53, 54, 57

Details:

• N [33] Additional rows are assumed to represent increasing complexity

• N [52] The presence of additional plates in a shell field is considered to represent the derived state; this character might be better expressed as two: 52a, multiple plates in transverse shell field; 52b, number of plates in transverse shell field - inapplicable if 52a = FALSE

63 taxa, 150 characters, of which 44 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 17, 18, 19, 20, 21, 22, 24, 25, 26, 27, 33, 37, 43, 44, 45, 49, 51, 52, 55, 59, 64, 71, 73, 78, 98, 105, 108, 110, 114, 115, 117, 123, 124, 125

• Neomorphic: 6, 8, 9, 23, 29, 31, 36, 122, 127 ; with state 1 denoting absence: 126

Details:

• N [122] Differentiation taken as derived

55 taxa, 88 characters, of which 27 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 6, 7, 8, 9, 10, 11, 12, 14, 15, 20, 21, 22, 23, 47

• Neomorphic: 5, 13, 16, 17, 18, 19, 46, 48, 59, 62, 63, 64, 77

Details:

• X [1] Degree to which the introvert can be invaginated: no introvert, or introvert not invaginable at all (0), partially invaginable (i.e., part of zone I) (1), completely invaginable into the trunk (i.e., to the base of zone I) (2).

• X [2] Anterior branched frontal appendages: absent (0), present (1).

• X [3] Scalids (nonspecific and sensu lato, including both scalids and presumed scalid derivatives): absent (0), present and composed exclusively of cuticle (1), present and with cuticle limited to a thin outer covering (2). Schmidt-Rhaesa (1998) has demonstrated that, in comparison to the scalids of Scalidophora (Kinorhyncha +Loricifera+Priapulida), the scalids of nematoids (Nematoda+Nematomorpha) are composed exclusively of cuticle. The collapse and folding of the scalids in Markuelia indicates that they are hollow structures and, thus, not wholly composed of cuticle.

• X [4] Mouth cone: absent (0), present but non-inversible (1), present and inversible (2). The mouth cone is the eversible (though not necessarily inversible) upstanding anterior limit of the pharynx.

• N [5] Zone I: unarmed (0), armed (1).

• T [6] Arrangement of zone I armature: not in rows (0), discrete parallel longitudinal rows (1), rows aligned diagonal to the anterior-posterior axis of the animal (2).

• T [7] Radial arrangement of armature: pentaradial (0), hexaradial (1).

• T [8] Morphology of zone I armature: papillae (0), simple spines (1), hooks or spinose hooks (2), conical scalids (3), telescopiform scalids (4), curved scalids and dentoscalids (5), complex scalids (6), glandular scalids, trifid spines, sensory spines, and double, tentaculite scalids (7), scalids with pectinate hood (8)*, spinoscalids and clavoscalids (kinorhynchs and lorificerans) (9).

• T [9] Number of elements comprising the first three circlets and, hence, defining the number of longitudinal rows of elements on the introvert: none (0), < 20 (1), 25 (2), > 25 (3). In all extant priapulids there are eight elements in the most proximal circlet of the introvert, their number and position corresponding to innervation derived from the circumpharyngeal brain.

• T [10] Sequence of zone I elements: elements as a single series (all elements identical or with differing morphologies) (0), elements organized into two or more transverse bands or series, possibly with different element morphologies within each series, but the sequence of morphologies being comparable between subsequent series (1).

• T [11] Basal circlet of zone I armature separated from more anterior by a constriction (as in loriciferans) or by insertion of longitudinal or circular muscles (kinorhynchs): absent (0), present (1).

• T [12] Number of elements in the proximal circlet of zone II: none or zone unarmed (0), eight (1), more than eight (2).

• N [13] Elongate spines at the base of Zone II: absent (0), present (1).

• T [14] Number of circlets of zone III armature: none or unarmed (0)*, one to four (1), six to fourteen (2), sixteen or more (3).

• T [15] Morphology of proximal circlets of zone III armature (“teeth”): spines or papillae (0), multispinose (1), multispinose but massively reduced (2), hooks (3), conical with a fringe of spines (4), sclerotized trabeculae (5), pectinate (6), conical papillae terminating in a long spine (prickle) (7), oral stylets (8).

• N [16] Morphology of middle circlets of zone III armature (teeth): absent (0), spines (of any length) or papillae (1), multispinose (reduced or otherwise) (2), pectinate (3).

• N [17] Morphology of the distal circlets of zone III armature (teeth): absent (0), spines (of any length) (1), multispinose (reduced or otherwise) (2), pectinate (3).

• N [18] Number of elements in first circlet of pharyngeal armature (base of zone III): elements absent (0), first circlet of 5 elements (1), first circlet of 8-10 elements (2), first circlet of numerous elements (more than 10) (3).

• N [19] Number of proximal, pentagonal circlets in zone III of the proboscis: none (0), five (1), six (2), seven (3).

• T [20] Width of zone III relative to zone II: zone III less than twice the width of zone II (0), zone III equal to or greater than twice the width of zone II (1).

• T [21] Width of the distal portion of zone Ill: distal zone III parallel to proximal zone III or tapering gradually (0), distal zone III expanded into a bulb (1).

• T [22] Eversibility of zone Ill: zone III completely eversible (0), zone III incompletely eversible, but eversible beyond the proximal teeth (1), zone III normally eversible only as far as the proximal teeth (2).

• T [23] All zone III elements of approximately equal size (0), zone III elements decreasing regularly in size from the posterior to the anterior (anteriormost elements less than half the size of the posteriormost) (1).

• X [24] Armament of cervical region: none (0), cuticular plates (as in larvae/juveniles) (1).

• X [25] Aspect ratio of body length to width in adult: <10 (0), 10-20 (1), >20 (2).

• X [26] Paired, lateral, locomotory appendages: absent (0), present (1).

• X [27] Annulation: absent (0), homonomous (1), heteronomous (2). Budd (2003) has drawn comparison between the patterns of cuticular ornament in association with the trunk annulae in lobopods and palacoscolecids. However, although palaeoscolecids exhibit an alternating pattern of cuticular ornament (Conway Morris 1997) and although this varies from one body region to another, adjacent annulae are identical; they exhibit homomous annulation. This contrasts with the condition in lobopods where adjacent annulae vary both in terms of their axial length and their cuticular ornamentation.

• X [28] Number of trunk annuli: <7 (0), 7-11 (1), 30-60 (2), 61-120 (3), 140 or more (4).

• X [29] Trunk spines, fine spines, or setae in adult: absent (0), present (1).

• X [30] Trunk papillae in adult: absent (0), present and simple (1), extremely long and arranged into two discrete, longitudinal rows (2)*.

• X [31] Trunk sclerites in adult: absent (0), present and small to medium in size (1), present and greatly enlarged (2).

• X [32] Trunk sclerites: absent (0), in two longitudinal rows (1), in annular rings (2).

• X [33] Trunk tumuli: absent (0), present (1).

• X [34] Trunk tubuli: absent (0), present (1).

• X [35] Trunk tubercles of soft, coriaceous consistency: absent (0), present (1).

• X [36] Flosculi, N-flosculi or sensory spots: absent (0), flosculi present with between 7 and 11 petals (1), flosculi present and invariably with 8 petals (2).

• X [37] Clavulae: absent (0), present (1).

• X [38] Tube: Absent (0), tube composed of plant debris: absent (1), tube flattened, annulated, tapering and chitinous (2).

• X [39] Arc or ring of posterior spines or hooks (distinct from the bilaterally symmetrical structures present in Markuelia and many palaeoscolecids): absent (0), present (1).

• X [40] Posterior ring papillae: absent (0), present (1).

• X [41] Eversible bursa: absent (0), present (1).

• X [42] Posterior abdomen greatly extensible: no (0), yes (1).

• X [43] Position of the anus: anus terminal, whether within bursa or otherwise (0), anus in posterolateral or posteroventral surface of the abdomen (1).

• X [44] Posterior tubulae or setae: absent (0), present (1).

• X [45] Caudal appendage(s): absent (0), present but less than the length of the body (1), present and up to three times the lenth of the body (2). Harvey et al. (2010) coded a caudal appendage as present in Kerygmachela. However, other characters contingent on the presence of a caudal appendage are coded as inapplicable for Kerygmachela by these authors (as for taxa lacking a caudal appendage). We therefore code an appendage as absent in Kerygmachela.

• N [46] Division of caudal appendage(s) or tail: undivided (0), pseudosegmented (1).

• T [47] Caudal appendage(s): single and positioned terminally (0), single and positioned dorso-medially (1), bicaudal as in some species of Priapulopsis and Paratubiluchus (2).

• N [48] Surface of caudal appendage: smooth (0), vesiculae (1), large warts (2).

• X [49] Terminally posterior spines, hooks or cones of basal diameter >20% trunk diameter: absent (0), present and arranged in pairs about the sagittal plane (1). This definition of ‘spines, hooks or cones’ excludes the posterior body bifurcations of Aysheaia, Kerygmachela, tardigrades, some nematomorphs, some kinorhynchs and gastrotrichs. The paired, peri-anal structures in Halicryptus and Maccabeus are much smaller relative to body width.

• X [50] Posterior warts: absent (0), small (as in Halicryptus) (1), large (as in Priapulus) (2).

• X [51] Polythyridium: absent (0), present (1).

• X [52] Ventral nerve cord unpaired throughout its length: absent (0), present (1). Living priapulids possess unpaired ventral nerve cords, whereas gastrotrichs, onychophorans and loriciferans possess ventral nerve cords that are paired throughout their length, and the ventral nerve cords of nematomorphs and nematodes divide at points along their length (Schmidt-Rhaesa, 1998); the situation in kinorhynchs is unresolved (paired according to Kristensen and Higgins, 1991; unpaired according to Neuhaus 1994). The condition in Ottoia is common to extant priapulids (Conway Morris, 1977).

• X [53] Ventral nerve cords merge caudally: absent (0), present (1).

• X [54] Dorsal nerve cord unpaired: absent (0), present (1).

• X [55] Circumpharyngeal brain: absent (0), present (1).

• X [56] Brain with anterior posterior sequence of perikarya-neuropil-perikarya: absent (0), present (1). Lemburg (1999) recognized this as a synapomorphy of Introverta (Nematoda + Nematomorpha + Kinorhyncha + Loricifera + Priapulida).

• X [57] Apical part of the brain composed of perikarya: absent (0), present (1).

• X [58] Pharyngeal nervous system composed of numerous tooth ganglia connected by a diagonal nerve net: absent (0), present (1). Lemburg (1999) recognizes the presence of this character as a synapomorphy of (extant) Priapulida.

• N [59] Two rings of introvert retractors attached through the collar-shaped brain: absent (0), present (1). Proposed by Nielsen (2001) as a synapomorphy of kinorhynchs, loriciferans and extant priapulids.

• X [60] Developmental mode: direct (0), biphasic (1).

• X [61] Loricate stage: absent (0), present (1).

• N [62] Cuticle of the larva/juvenile dorso-ventrally flattened (at least in older stages), with six lateral plates in-folded and accordion like. Cuticle of the lorica thickened in dorsal and ventral plates (at least) with sculpture of four to six longitudinal rows of narrow, rectangular fields: absent (0), present (1).

• N [63] Larvae/juveniles with six long pharynx retractor muscles: absent (0), present (1). Lemburg (1999) recognised the presence of this character as a synapomorphy of (extant) Priapulida. However, it has since been demonstrated that the larvae of nematomorphs also possess six pharyngeal retractor muscles (Kristensen, 2003; Muller et al., 2004).

• N [64] Teeth of second circle of the larvae/juveniles with very small median denticle: absent (0), present (1). Lemburg (1999) recognizes the presence of this character as a synapomorphy of Megaintroverta (Priapulopsis + Acanthopriapulus + Priapulus).

• X [65] Division of the body into a distinct proboscis and abdomen in larva/juvenile: absent (0), present (1).

• X [66] Cloaca in both sexes: absent (0), present (1).

• X [67] Protonephridia: absent (0), present but do not flow into the gonoduct and are not integrated into the gonad (1), present and flow into the gonoduct and/or are integrated into the gonad (2). Protonephridia are considered an apomorphy of the Bilateria (Ax, 1996) and are present in gastrotrichs, kinorhynchs, loriciferans and extant priapulids, but absent, presumably secondarily, from onychophorans, nematodes and nematomorphs. Lemburg (1999) recognized state 2 as a synapomorphy of lorciferans and extant priapulids.

• X [68] Urogenital system attached to the body wall by a ligament: absent (0), present (1). Lemburg (1999) recognized this as a synapomorphy of lorciferans and extant priapulids.

• X [69] Spermatozoa with a flagellum: absent (0), present (1). The presence of a flagellum in spermatozoa is a metazoan symplesiomorphy, but a flagellum is lacking from the spermatozoa of nematodes and nematomorphs (Schmidt-Rhaesa 1998).

• X [70] Genital field extremely simplified: no (0), yes (1).

• X [71] Copulatory organs: absent (0), present (1).

• X [72] Perigenital setae: absent or reduced (0), prominent (1).

• X [73] Cuticle predominantly containing collagen: absent (0), present (1).

• X [74] Distribution of chitin in the cuticle: none (0)*, predominantly within the middle cuticle layer (exocuticle) (1), predominantly within the lowermost cuticle layer (endocuticle) (2).

• X [75] Cuticle with homogeneous layer at surface/beneath epicuticle: absent (0), present (1).

• X [76] Cuticle with middle layer of distinct composition: absent (0), present (1).

• N [77] Cuticle with radially striated/vertical canal middle layer: absent (0), present (1).

• X [78] Cross-wise fibres in cuticle: absent (0), present (1). (Reported in Meiopriapulus by Storch et al. 1989).

• X [79] Large helical fibres in the cuticle: absent (0), present and composed of platy subunits (1), present and composed of unpaired fibrils (2), present and composed of paired, wound fibrils (3).

• X [80] Cuticle surface with ornament of tessellating polygons: absent (0), present (1).

• X [81] Division of the body into a distinct proboscis and abdomen in adult: absent (0), present (1).

• X [82] Terminal mouth: absent (0), present (1).

• X [83] Percentage of animal length taken up by the introvert: None (no introvert) (0), 2% (as in Halicryptus higginsi) (1)*, between 2% and 30% (2), 30 to 50% (3). Lemburg (1999) recognizes a very large introvert (state 3) as a synapomorphy of Megaintroverta (Priapulopsis + Acanthopriapulus + Priapulus).

• X [84] Voluminous primary body cavity: absent (0), present (1). Lemburg (1999) recognizes the presence of this character as a synapomorphy of (extant) Priapulida.

• X [85] Movement by peristaltic movement of the pharynx/introvert: absent (0), present (1).

• X [86] Pharyngeal lumina: round (0), triradiate (1).

• X [87] Circular body musculature: absent (0), present (1). Circular body musculature is present in all taxa of nemathelminth grade except nematodes and nematomorphs, and its absence has been considered both secondary and a synapomorphy of Nematoida (Schmidt-Rhaesa 1998).

• X [88] Nucleation of “peritoneal” membrane: membrane without nuclei or simply with amoebocytes in association with the surface (0), membrane containing scattered nuclei (1).

61 taxa, 165 characters, of which 70 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 2, 10, 13, 24, 26, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 44, 45, 46, 47, 48, 49, 61, 65, 72, 74, 80, 85, 94, 95, 96, 99, 101, 102, 105, 106, 115, 116, 117, 118, 121, 123, 129, 133, 134, 137, 138, 139, 140, 142, 143, 145, 146

• Neomorphic: 9, 148 ; with state 1 denoting absence: 1, 3, 4, 5, 50, 51, 73, 83, 84, 91, 100, 103, 113, 119

Details:

• I? [1] Cervical_groove / present absent,

• T [2] ‘Mandibular (genal or cheek) groove’ / ‘smoothly indented, or with acute indentation’ absent,

• I? [3] Mandibular_notch / present absent,

• I? [4] Clypeal_notch / present absent,

• I? [5] Antennal_notch / present absent,

• X [6] ‘Pereonites 2-7 in dorsal view (relative lengths)’ / longer_than_wide wider_than_long anteriorly_longer_than_wide_decreasing_posteriorly_to_wider_than_long,

• X [7] ‘Coxal articulation of pereonites 2-4’ / free nearly_fused_or_fused,

• X [8] ‘Coxal articulation of pereonites 5-7’ / free nearly_fused_or_fused,

• N [9] Sternal_processes_on_sternites / present absent,

• T [10] ‘Typhlosole (form in cross-section)’ / ‘well developed, ventral invagination forming double spiral in cross section’ ‘intermediate, ventral invagination forming Y shape in cross section’ ’minimal, ventral invagination forming laminar projection or inverted u shape i

• X [11] ‘Pleonites in lateral view (depth of ventrolateral plates relative to pereonites)’ / ‘much deeper than pereonites, with large pleurae, basal region of pleopods not visible’ ’having depth approximating depth of pereonites, with small pleurae, basal region of pleopods

• X [12] Pleonites_with_pleonite_1_pleura / ‘distinctly shallower than pleurae of pleonites 2-5’ ‘near depth of pleurae of pleonites 2-5’,

• T [13] Pleonite_5_ventral_margin_shape / ‘subparallel to dorsal margin along length, not sinuate posteriorly’ sinuate,

• X [14] Pleotelson_form / ‘shallow (lateral fields sloping)’ ‘vaulted (lateral fields vertical)’,

• X [15] Pleotelson_dorsal_surface_shape_in_lateral_view / evenly_curving strongly_inflected_ventrally,

• X [16] Pleotelson_dorsal_surface_setae / lacking_simple_setae sparsely_covered_with_fine_setae covered_with_abundant_curled_simple_setae covered_with_abundant_elongate_setae densely_covered_with_cuticular_hairs_and_setae with_abundant_robust_setae,

• X [17] Pleotelson_dorsal_surface_median_ridge / absent present,

• X [18] Pleotelson_dorsal_surface_lateral_ridges / continuous_with_posterior_apex absent,

• X [19] Pleotelson_lateral_length_versus_depth / less_than_depth_or_subequal_to_depth greater_than_depth,

• X [20] Pleotelson_ventral_surface_anterior_to_uropods / strongly_concave ‘flattened, only slightly concave’,

• X [21] Pleotelson_ventral_margin_anterior_to_uropods_robust_setae / with_robust_setae with_distally_denticulate_setae_in_one_row with_distally_denticulate_robust_setae_in_two_rows with_fine_setae_only,

• X [22] Pleotelson_ventral_margin_relative_lengths_of_robust_setae_anterior_to_uropods / posterior_seta_longer_than_anterior_adjacent_setae posterior_seta_length_subequal_to_anterior_adjacent_setae posterior_seta_shorter_than_anterior_adjacent_setae,

• X [23] Pleotelson_postanal_ventral_surface / ‘present (apical margin and anus separated by cuticular surface)’ ‘absent (anus adjacent to apical margin)’,

• T [24] Pleotelson_postanal_ventral_surface_ornamentation / ‘unelaborated, or with fine setae on ridge’ ‘with robust setae (not on ridge)’ with_robust_setae_on_ridge with_groove_forming_cleft_in_posterior_apex,

• X [25] Pleotelson_lateral_uropodal_ridge / terminating_at_pleotelson_margin_above_uropods curving_strongly_and_extending_posteriorly_from_uropods absent,

• T [26] Pleotelson_lateral_uropodal_ridge_setae / with_row_of_robust_setae with_1_robust_seta with_fine_simple_setae lacking_setae,

• X [27] ‘Pleotelson posterolateral margin, differentiation from apex’ / uninterrupted with_distinct_inflection_differentiating_apex,

• X [28] Pleotelson_posterolateral_margin_shape / forming_rounded_process_with_dorsally_projecting_setae crenate forming_vertical_plate forming_rounded_lobe unelaborated,

• T [29] ‘Pleotelson posterolateral margin (shape if lobe-like)’ / curving_dorsally depressed_ventrally_and_posteriorly_angular,

• T [30] ‘Pleotelson posterolateral margin robust setae (if plates or rounded lobes presen’ / without_robust_setae with_one_position_with_robust_setae with_two_positions_with_robust_setae_with_intermediate_setae with_two_positions_with_robust_setae_without_intermediate_setae

• T [31] ‘Pleotelson posterolateral margin dorsal robust setae (if plates or lobes present’ / with_one_robust_seta ‘with multiple robust setae, ventrolateral seta’ ‘with multiple robust setae, ventrolateral setae subequal to dorsomedial setae’,

• T [32] ‘Pleotelson posterolateral margin submarginal seta near inflection point (if plat’ / present absent,

• T [33] ‘Pleotelson posterolateral margin dorsolateral surface robust setae (if lobe-like’ / absent adjacent_to_margin set_anteriorly_from_margin,

• T [34] ‘Pleotelson posterior apex shape in dorsal view (if postanal ventral surface pres’ / projecting indented transverse,

• T [35] ‘Pleotelson posterior apex (flattened or not)’ / ‘free (not flattened)’ strongly_reflexed_and_flattened_against_dorsal_surface,

• T [36] ‘Pleotelson posterior apex projections (if postanal ventral surface absent)’ / with_small_rounded_tubercle without_projections,

• T [37] ‘Pleotelson posterior apex length in lateral view(if projecting)’ / ‘very long (> 0.295 pleotelson length)’ ‘long (0.120-0.295 pleotelson length)’ ‘short (< 0.119 pleotelson length)’,

• T [38] ‘Pleotelson posterior apex shape in dorsal view (if protruding freely)’ / elongate broadly_rounded quadrate,

• T [39] Pleotelson_posterior_apex / visible_in_lateral_view obscured_by_posterolateral_margin_in_lateral_view,

• T [40] Pleotelson_posterior_apex_robust_setae / absent with_1_pair_of_robust_setae with_2_pairs_of_robust_setae with_3_pairs_of_robust_setae with_4_pairs_of_robust_setae,

• X [41] ‘Antennula articles (in male)’ / more_than_6_articles 6_or_fewer_articles,

• X [42] Antennula_article_3_rudimentary_second_flagellum / present absent,

• X [43] ‘Antennula article 4 length (relative to article 3)’ / shorter ‘subequal (ratio 0.95-1.05)’ longer,

• T [44] ‘Antennula terminal article shape (taxa with more 6 articles in male)’ / tubular globular ‘distally oblique, with 2 or more groups of aesthetascs’ conical ‘tiny, vestigial’,

• T [45] ‘Antennula terminal article length relative to penultimate article (taxa more tha’ / shorter subequal longer,

• T [46] Antennula_penultimate_article_length / longer_than_any_other_article ‘length approximately sub-equal to, or less than length of other articles’,

• T [47] ‘Antennula articles 5-6 shape (taxa with 6 or fewer articles in male)’ / inflated_and_bulbous width_subequal_to_article_4 article_5_only_inflated_and_bulbous,

• T [48] ‘Antennula article 5 length relative to article 6 (taxa with 6 or fewer articles’ / subequal_or_less greater article_6_not_expressed,

• T [49] ‘Antennula penultimate and ante-penultimate articles shape in cross section’ / flattened ‘oval, or circular’,

• I [50] ‘Antenna flagellum proximal articles dense cover of cuticular hairs (in male)’ / present absent,

• I [51] ‘Antenna flagellum proximal articles dense cover of cuticular hairs (in female)’ / present absent,

• X [52] Antenna_propodal_article_3_scale / well_developed rudimentary_but_articulated absent,

• X [53] Antenna_article_5_length_relative_to_article_4 / shorter_or_subequal longer,

• X [54] Antenna_article_6_length_relative_to_articles_4_and_5_combined / shorter_or_subequal longer,

• X [55] Mandible_palp_third_article_shape / ‘crescent-like’ relatively_linear,

• X [56] ‘Mandible palp third article setae on medial-distal margins’ / with_fewer_than_5_setae with_more_than_5_setae,

• X [57] Mandible_palp_3rd_article_coarsely_spinulate_setae / absent coarsely_spinulate_setae_with_other_setae coarsely_spinulate_setae_present_only,

• X [58] Mandible_palp_third_article_medial_surface / naked covered_in_dense_cuticular_hairs covered_in_cuticular_combs,

• X [59] Mandible_incisor_processes_width_relative_to_thickness / broad thin,

• X [60] Mandible_right_lacinia_mobilis_shape / ‘large, distinctly separated’ ‘indistinct, spine-like’ absent,

• T [61] Mandible_right_lacinia_mobilis_dentate_plates / two_dentate_plates single_dentate_plate,

• X [62] Mandible_spine_row_bifurcate_spines / present absent,

• X [63] Mandible_spine_row_general_shape_between_incisor_and_molar / in_smooth_low_arc on_projecting on_linear_pedunculate_projection on_round_pedunculate_projection on_elongate_penicil,

• X [64] Mandible_spine_rows_shape_of_spine_ridge_in_ventral_view / distal_margin_in_line_with_proximal_margin distal_margin_protruding_in_relative_to_proximal_margin ‘forming strongly convex arc, protruding medially’,

• T [65] Mandible_spine_rows_shape_of_spine_ridge_in_medial_view / basal_insertions_in_line_between_incisor_and_molar_processes basal_insertions_crossing_dorsally_and_then_abruptly_angling_posteriorly,

• X [66] Mandible_right_spine_row_first_spine_relative_to_remaining_spines / separated not_separated,

• X [67] Maxillula_medial_lobe_width / less_than_or_subequal_to_lateral_lobe greater_than_lateral_lobe,

• X [68] Maxillula_medial_lobe_pappose_setae / up_to_three_setae four_setae five_setae six_setae_or_more,

• X [69] Maxillula_medial_lobe_accessory_setae / one_or_two more_than_two absent,

• X [70] Maxillula_lateral_lobe_distal_margin_denticulate_robust_setae / with_one_or_more absent,

• X [71] Maxillula_lateral_lobe_distal_margin_smooth_robust_setae / up_to_nineteen twenty_or_more,

• T [72] Maxillula_lateral_lobe_ventral_face_setae_position / widely_spaced closely_spaced,

• I [73] ‘(Maxillula) lateral lobe ventral face plumose seta’ / present absent,

• T [74] Maxilla_outer_lateral_lobe_distal_margin_setal_row / approximately_linear_and_transverse_to_lobe_axis curving_and_extending_proximally_along_medial_margin ‘with two angles - transverse to lateral margin and oblique on medial margin’,

• X [75] ‘Pereopod I dactylus ventrodistal margin cuticular fringe (in male)’ / with_multiple_rows_of_minute_sharp_spines with_row_of_sharp_spines ‘with row of thin scale-like spines’ ‘with cuticular fringe (minute cuticular hairs)’ cuticular_fringe_absent,

• X [76] ‘Pereopod I dactylus ventrodistal margin cuticular fringe (in female)’ / with_multiple_rows_of_minute_sharp_spines with_row_of_sharp_spines ‘with row of thin scale-like spines’ cuticular_fringe_absent,

• X [77] Pereopod_I_dactylus_accessory_claws / several one_only absent,

• X [78] Pereopod_I_dactylus_distal_accessory_spines / several one_only absent,

• X [79] ‘Pereopod I propodus dorsal margin setae (in male)’ / setae_present_in_several_groups_between_proximal_and_distal_margin setae_confined_to_single_group_at_distal_margin,

• T [80] ‘Pereopod I propodus dorsal margin proximal region (in male)’ / protruding_to_distodorsal_margin_of_ischium protruding_to_distodorsal_margin_of_carpus not_protruding,

• X [81] ‘Pereopod I propodus dorsal margin proximal region (in female )’ / protruding_to_distodorsal_margin_of_ischium protruding_to_distodorsal_margin_of_carpus not_protruding,

• X [82] ‘Pereopod I propodal palm shape (in male)’ / concave convex_to_straight,

• I [83] ‘Pereopod I propodal palm proximal simple spine (in male)’ / present absent,

• I [84] ‘Pereopod I propodal palm median simple spine (in male)’ / present absent,

• T [85] ‘Pereopod I propodal palm distal simple spines (in male)’ / one more_than_one absent,

• X [86] ‘Pereopod I propodal palm medial rugose cuticular pad (in male)’ / present absent,

• X [87] ‘Pereopod I propodal palm stout denticulate setae (in male)’ / serrate bifid absent,

• X [88] ‘Pereopod I propodal palm stout robust setae (in male)’ / conical basally_inflated absent,

• X [89] ‘Pereopod I propodal palm setal ridge (in male)’ / absent present,

• X [90] ‘Pereopod I propodal palm shape (in female)’ / concave convex straight,

• I [91] ‘Pereopod I propodal palm midpoint simple spine (in female)’ / present absent,

• X [92] ‘Pereopod I propodal palm stout denticulate setae (in female)’ / serrate bifid absent,

• X [93] ‘Pereopod I propodal palm stout robust simple setae (in female)’ / conical basally_inflated absent,

• T [94] ‘Pereopod I merus distodorsal margin in anterodorsal view (in male)’ / ‘shelf-like and U-shaped’ ‘spine-like and pointed’,

• T [95] ‘Pereopod I merus distodorsal margin projection in anterodorsal view (in female)’ / ‘shelf-like and U-shaped’ ‘spine-like and pointed’,

• T [96] ‘Pereopod I merus distodorsal margin setal form (in male)’ / with_more_than_2_elongate_simple_setae ‘with 1-2 robust simple setae’ ‘with 1-2 elongate simple setae’,

• X [97] ‘Pereopods II-III propodus articular plate’ / present absent,

• X [98] ‘Pereopod II basis dorsal margin proximal group of setae (in male)’ / robust_setae elongate_fine_setae without_distinct_group_of_large_setae,

• T [99] ‘Pereopod II basis dorsal margin proximal group of setae (in male)’ / in_1_row ‘in 2 rows (rarely more)’,

• I [100] ‘Pereopod II basis dorsal margin projection (in male)’ / present absent,

• T [101] ‘Pereopod II basis dorsal margin proximal group of setae (in female)’ / robust_setae elongate fine setae proximally without_distinct_group_of_large_setae,

• T [102] ‘Pereopod II basis dorsal margin proximal group of setae number of rows (in femal’ / in_1_row ‘in 2 rows (rarely more)’,

• I [103] ‘Pereopod II basis dorsal margin projection (in female)’ / present absent,

• X [104] ‘Pereopod IV prehensility (sexual dimorphism, adult male form)’ / prehensile ‘similar to female, not prehensile’,

• T [105] ‘Pereopod IV subchelate major hinges (in male if dimorphic)’ / on_dactylus_and_propodus on_propodus_and_carpus,

• T [106] ‘Pereopod IV dactylus length (in male)’ / shorter_than_or_subequal_to_propodal_palm longer_than_propodal_palm,

• X [107] Pereopod_IV_propodus_articular_plate / present absent,

• X [108] ‘Pereopods V-VII propodus articular plate’ / present absent,

• X [109] ‘Pereopods V-VII basis dorsal ridge shape’ / rounded_or_angular_on_all forming_distinct_plate_at_least_on_pereopod_VII,

• X [110] ‘Pereopods V-VII basis dorsal margin’ / with_robust_setae with_elongate_fine_setae without_large_setae,

• X [111] ‘Pereopods V-VII basis lateral face central longitudinal ridge’ / present absent,

• X [112] ‘Pereopods V-VII basis lateral face ventral longitudinal ridge’ / present absent,

• I [113] ‘Pereopods V-VII basis lateral face ventral longitudinal ridge’ / with_fine_setae setae_absent,

• X [114] Pereopod_VII_ischium_dorsal_ridge_flange_width / greater_than_or_subequal_to_shaft_width less_than_shaft_width flange_absent,

• T [115] Pereopod_VII_basis_dorsal_ridge_distal_margin_shape / indented rounded,

• T [116] Penes_form / straight curved_posteriorly,

• T [117] Penes_extent_medially / near_or_reaching_midline past_midline_and_onto_first_pleonite,

• T [118] Penes_shaft_cuticle / smooth with_cuticular_hairs denticulate,

• I [119] Penes_setae / on_shaft on_tip absent,

• X [120] Penes_distal_tip_shape / rounded pointed flattened truncate,

• T [121] ‘Pleopod exopods II-V proximal article distolateral lobes length relative to dist’ / shorter_than_distal_article subequal_or_longer_than_distal_article,

• X [122] ‘Pleopod exopods I-V lateral proximal lobes’ / ‘on I-V’ ‘on II-V’ absent_on_II absent,

• T [123] ‘Pleopod exopods I-V medial proximal lobes’ / ‘on I-V’ ‘on II-V’ ‘on III-V’ absent,

• X [124] ‘Pleopod endopods marginal setation (in male)’ / ‘endopods I-V without setae’ endopod_I_only_with_setae ‘endopods I-V with setae’,

• X [125] Pleopod_protopods_medial_margins / ‘I-V with epipods or small projections’ ‘II-V with epipods or small projections’ ‘III-V with epipods’ ‘I-V without epipods’,

• X [126] ‘Pleopod I-V protopods medial margin coupling hooks (in male)’ / ‘present (on at least some pleopods)’ absent,

• X [127] ‘Pleopod protopods medial margin setae (in male)’ / with_plumose_setae with_minutely_serrate_or_simple_setae setae_absent,

• X [128] ‘Pleopod protopods medial margin coupling hooks (in female)’ / present absent,

• T [129] ‘Pleopod protopods medial margin pleopod I coupling hook numbers (in female)’ / one_only with_multiple_coupling_hooks,

• X [130] Pleopod_protopod_I_lateral_epipod / ‘lobe-like’ linear absent,

• X [131] Pleopod_protopod_II_lateral_epipod / ‘lobe-like’ linear absent,

• X [132] ‘Pleopod protopods lateral epipods III-V’ / ‘lobe-like’ absent,

• T [133] Male_pleopod_I_exopod_lateral_margin_shape / rounded broadly_angular concave convoluted,

• T [134] Male_pleopod_I_exopod_dorsal_surface / with_setae lacking_setae,

• X [135] Male_pleopod_I_exopod_ventral_surface_shape / flat curving_anteriorly,

• X [136] Male_pleopod_I_protopod_shape / longer_than_wide width_subequal_length significantly_wider_than_long,

• T [137] ‘Male pleopod II endopod appendix masculina (stylet) proximal shape ventrallly’ / broadly_concave ‘rod-like, convex’ ‘rod-like, indented’,

• T [138] Male_pleopod_II_endopod_stylet_channel / tube_along_distal_length not_forming_tube,

• T [139] Male_pleopod_II_endopod_stylet_tip_shape / truncateor_broadly_rounded acutely_rounded conical ‘sharply pointed and spine-like’,

• T [140] Male_pleopod_II_endopod_stylet_tip_denticles / smooth_or_with_minute_rounded_denticles with_large_subterminal_denticles,

• X [141] Male_pleopod_II_endopod_stylet_marginal_setation / multiple_setae 1_seta setae_absent,

• T [142] Male_pleopod_II_endopod_stylet_length_relative_to_exopod / near_to_distal_margin not_extending_beyond_half_length,

• T [143] Uropod_protopod_extension_posteriorly_in_lateral_view / beyond_or_subequal_to_pleotelson_apex not_reaching_pleotelson_apex,

• X [144] Uropod_protopod_dorsomedial_ridge_shape / ‘plate-like’ ‘spur-like’ rounded_projection,

• T [145] Uropod_protopod_dorsomedial_ridge_shape / parallel_to_ventral_margin laterally_concave_at_midpoint,

• T [146] Uropod_protopod_dorsomedial_ridge_margin_setal_form / robust_spinose_distally robust_simple fine_simple absent,

• X [147] Uropod_protopod_distoventral_margin_robust_setal_number / absent one two three four_or_more,

• N [148] Uropod_protopod_distoventral_margin_robust_spinose_setae / absent present,

• X [149] Uropod_rami_distal_tips_shape / pointed rounded,

• X [150] ‘Uropod rami cross-sectional shape’ / round flattened_dorsally_and_ventrally flattened_on_dorsal_surface_only ‘exopod round, endopod flattened’,

• X [151] Uropod_endopod_length_relative_to_protopod / shorter subequal longer,

• X [152] Uropod_endopod_length_relative_to_exopod / subequal_or_longer distinctly_shorter,

• X [153] Uropod_endopod_shape / straight_or_curving_dorsally curving_medially,

• X [154] Uropod_endopod_dorsal_margin_robust_setae_placement / all_along_length midlength proximally absent,

• X [155] Uropod_endopod_spine_on_dorsal_margin / present absent,

• X [156] Uropod_endopod_proximoventral_margin_shape_in_medial_view / sinuate convex,

• X [157] ‘Uropod exopod robust setae on dorsal margin (in male)’ / one_or_more absent,

• X [158] ‘Uropod exopod robust setae on dorsal margin (in female)’ / one_or_more absent,

• X [159] Pereopod_II_basis_proximal_dorsal_ridge / absent present,

• X [160] Pereopodal_exopods / present absent,

• X [161] Short_carapace / present absent,

• X [162] Heart_position / thoracic ‘pleonal-thoracic’,

• X [163] Uropodal_articles / either_multiarticulate uniarticulate,

• X [164] Telson / free fused_to_pleonite_6,

• X [165] Antennular_second_flagellum / multiarticulate tiny_or_absent,

37 taxa, 105 characters, of which 14 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 11, 12, 16, 27, 36, 45, 91

• Neomorphic: 18, 29, 30, 31, 32, 61, 62

Details:

• N [62] Same length presumed ancestral

74 taxa, 213 characters, of which 150 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 9, 11, 12, 13, 16, 17, 18, 21, 23, 24, 26, 28, 30, 31, 32, 34, 35, 38, 41, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52, 54, 55, 56, 58, 59, 60, 61, 63, 64, 65, 66, 67, 70, 72, 73, 74, 77, 78, 79, 80, 81, 82, 87, 88, 89, 90, 91, 92, 93, 94, 103, 104, 105, 106, 107, 112, 113, 114, 115, 118, 119, 120, 121, 123, 124, 125, 126, 127, 129, 136, 137, 138, 139, 144, 145, 146, 147, 148, 149, 150, 151, 152, 153, 154, 155, 156, 157, 158, 159, 160, 161, 162, 163, 164, 165, 166, 167, 168, 169, 174, 175, 176, 177, 178, 179, 180, 181, 182, 183, 184, 189, 190, 191, 193, 194, 195, 196, 197, 198, 199, 200, 201, 202, 205, 206, 207, 208, 209

• Neomorphic: 7, 10, 22, 25, 29, 33, 42, 53, 71, 108, 109, 192, 213

75 taxa, 253 characters, of which 179 contain inapplicable codings.

Download: Manually revised matrices | Inapplicable | Extra State | Ambiguous

Inapplicable characters:

• Transformational: 5, 9, 14, 16, 19, 21, 22, 23, 25, 26, 29, 30, 32, 34, 36, 41, 44, 47, 53, 70, 71, 78, 80, 87, 90, 105, 106, 107, 111, 112, 118, 119, 125, 129, 136, 143, 148, 149, 150, 152, 153, 159, 160, 161, 162, 164, 167, 168, 169, 170, 172, 174, 175, 177, 178, 180, 181, 182, 183, 184, 185, 186, 187, 188, 189, 190, 192, 193, 194, 198, 200, 201, 203, 205, 209, 210, 211, 212, 215, 217, 218, 227, 228, 237, 238, 240, 243, 245, 246, 247, 250

• Neomorphic: 7, 10, 11, 12, 13, 24, 28, 31, 33, 37, 38, 39, 40, 42, 43, 45, 46, 48, 49, 50, 51, 52, 62, 77, 79, 99, 104, 108, 109, 110, 113, 114, 115, 116, 117, 120, 122, 123, 124, 127, 128, 131, 132, 133, 134, 139, 140, 141, 144, 145, 146, 147, 151, 154, 155, 156, 157, 163, 165, 171, 173, 176, 179, 195, 196, 197, 199, 202, 204, 206, 207, 208, 213, 214, 216, 224, 229, 235, 236, 239, 241, 242, 244, 248, 249, 251, 252, 253

Details:

• N [195] Loss assumed secondary