We took the phylogenetic analysis of pterosaur relationships by Hone et al. (2020) and modified it such that the holotype of Luchibang xingzhe and the now distinct postcranial material were coded as separate OTUs (see below). As in Hone et al. (2020), the OTU for Linlongopterus jennyae, which was pruned in the final analyses of Kellner et al. (2019) due to its status as a “wild card taxon,” was retained within our analysis for the sake of thorough sampling. Following the addition of these changes, we performed a phylogenetic analysis with the software TNT v 1.5 (Goloboff et al., 2018) using the TBR heuristic searches performed using maximum parsimony. Following Kellner et al. (2019), the search for the most parsimonious trees (MPTs) was conducted via Traditional Search (TBR swapping algorithm), 10,000 replicates, random seed, 20 trees to save per replication, and collapsing trees after search.
Changes made to the coding of the original Hone et al., 2020 analysis for revision are as follows:
3. Changed (1) to (?) as we don’t have enough to show a nasoantorbital fenestra is present.
22. Changed (0) to (?) as we don’t have enough to confidently say that the dorsal margin of the skull is entirely “nearly straight”.
42. Changed (0) to (?) because the absence of a posterior ventral expansion on the maxilla cannot be demonstrated with this portion of the skull.
43. Changed (0) to (?) because the absence of maxilla-nasal contact cannot be demonstrated.
44. Changed (-) to (?) for the reasoning in [43].
76. Changed (1) to (?) because the ratio cannot be determined with this fragment of mandible.
Removed all coding between 106-150 (?1?11????????102000?201??40??1??????212222120) as it is considered a separate specimen. This coding was moved to a new OTU, “Luchibang_postcrania”, with 1-105 coded as (?) because these cannot be coded for this specimen.
Postcranial OTU Changes
119. Changed (1) (“stout, with constructed shaft”) to (?) as this cannot be assessed.
124. Changed (0) to (1) as it is present.
128. Changed (1) to (0).
131. Changed (4) (“enlarged, warped”) to (5) (“long, proximally placed, curving ventrally”); there is no warping to the deltopectoral crest, and it best matches the condition of state (5).
132. Changed (0) to (1,2).
142. Changed (2) to (3).
146. Changed (2) to (3).
Consistency index: 0.630
Retention index: 0.872
Matrix:
xread
150 64
Ornithosuchus_woodwardi 00000020000010001000000100000-00000--00--0100---000000000000--0-0000000000?0000000--00-000000000000000000000000000000000000000000000000000000000000000
Herrerasaurus_ischigualastensis 00001000000000001000000100000-00000--00--0100---000000000000--0-000000000000000000--00-000000000000000000000000000000000000000000000000000000000000020
Scleromochlus_taylori 00001000000000001000020100000-00000--00--0100---00001?020??0--0-00????0000000?0000--00-00?000?000??000000???00??0000??0?000???00??0???0???00?0000000??
Anurognathus_ammoni 00000010000000001100120100000-00100--00--0110---000110010??0--0-000000000100000000--00-001000100100000000???0???01000?0???0??0110?100000??1001122?0011
Rhamphorhynchus_muensteri 11001000000110010100001300000-00000--00--0100---000000021000--0-000000001002012010--00-00100010000001000000000000010000010001111002000100?100122221012
Darwinopterus_linglongtaensis 1011000000011??????0001100000-0000?20000000-1010100010021?00--0-00?00?00????010000--00-0010001?1000000000???10??001?0?00?000?1110?100?10??100311111022
Darwinopterus_robustodens 101100?00?011??????0001100000-0000?20000000-1010?00010021?00--0-00?00?00????010000--00-0010001?1000000000???1???001?0?00??0??111??100?10??10031111102?
Pterodactylus_antiquus 1010000000011??0???0001100000-00000--00--00-1001000010031?00--0-10?00000???2010000--00-0010001010000000000002110010000001000121100500?100?100312222020
Dsungaripterus_weii 10100001011110411200001300000-0100111000010-?---0000100211012211011101001012012000--11-12--011000000000001111100110?000???0?2210??5?001001??13122?2120
Noripterus_parvus 1010000101111??????0001300000-0100111000010-?---000010021?01221101?10?00?0?2012000--?1-12--01100000000000?11??0??????????????????????0?001????????????
Keresdrakon_vilsoni 101????????????????00?1300000-?00?0--00--?0-???????????2????????????0000???301200101?3-------------------?111?00?????00020011?0?01520?1???????????????
Tupuxuara_leonardii 10110020020210511210001300100-0000134000000-?????0011002210127110110030010130120011313-------------------11111001???00002001?2100152011001?01313??2???
Thalassodromeus_sethi 1011002002021?611210001300100-0000134000000-1110000110022101271101100300101?0120011313-------------------?????????????????????????????????????????????
Caupedactylus_ybaka 1011002002021?511210021310100-0000134000000-?????00?210221?12711011?000010130120011013-------------------???????????00002001????015200????????????????
Aymberedactylus_cearensis ???????????????????00??31??????????????????????????????????????????????????20120011?13-------------------?????????????????????????????????????????????
Tupandactylus_imperator 1011002002021??????102131?100-0000133100000-??????0110?22?01221101????0????201210112?3-------------------?????????????????????????????????????????????
Caiuajara_dobruski 1011002002021??????1021311100-?000133100000-???????1?00220?1221101101001??120121011213-------------------?1111000?0?00002?01?21?11520?1??120??????????
Europejara_olcadesorum ??????2???021???????0????????????????????0?????????????220?12???0???????1?1201?10112?3-------------------?????????????????????????????????????????????
Tapejara_wellnhoferi 101100200202102????1021311100-0000133000000-?????1011002200122110110100110120121011213-------------------?1111000?0?00002001?210115200100120?31???2120
Sinopterus_dongi 1011002002021??????1021311000-1000133000000-1100010110022?11211101?0??01???20121011013-------------------?1?11000???0000200??211?152??100?20?3122321??
Huaxiapterus_corollatus 101????????????????102131?000-1000133010000-??????????0???????????????01????01210110?3-------------------?1?11??0???000???0??211??5??????????3132321??
Huaxiapterus_benxiensis 1011002002021??????102131?000-1000133010000-???????110022??1211101????01???201210110?3-------------------?1?11??0????????????2????????????20?3122?2120
Eopteranodon_lii 101??0???2?????????1021311000-100013300??00-????????????????????????????????012001???3-------------------????????????????????21???5??????????3?22?2???
Chaoyangopterus_zhangi 101??1?0???????????001130?000-?00????00--00-???????????????1??????????00????012000--?3-------------------?102110????000???10?200??????100?20?3132?21?0
Shenzhoupterus_chaoyangensis 1010012002021??????001130?000-0001155000000-?????0??2???2?01241101????00???2012000--?3-------------------?1021??????0?0???1??200??5???100????3122321??
Jidapterus_edentus_ 101????????????????0011300000-?001???00--00-??????????????????????????0????3012000--13-------------------?1?2110??????0???1??200??5?0?100????312232120
Azhdarcho_lancicollis ???????????????????00??3?000???0?????0?--?0-?????????????1??????????0?0??????????????3-------------------11031221???????????????015?0?????????????????
Quetzalcoatlus_sp. 10100020020????????0001300000-?00014?00--00-0---000010??210?????????000010?3012000--13-------------------11031221?0?00002010?20?0152001001???4132?2?20
Zhejiangopterus_linhaiensis 10100020020?1??????0001300000-00000------00-0---0000100?2?00--0-01??0?00????012000--?3-------------------1??31221?0?000???10?200??5???100?20?413??2???
Nyctosaurus_gracilis 10101000000110101100012300000-00000------00-?????00011021100--1200?000001013012000--13-------------------11?110111000?00100013110130?011122123222?20??
Muzquizopteryx_coahuilensis ?01010?000011???????0?????????0?00???????00-?????0001?021?00--1200????0??????????0???3-------------------???11011????1????????11??30??111?21???????1??
Pteranodon_longiceps 10100020000111302310012300000-00000------00-112?0000110231012311001000001013012000--03-------------------111110111011101200012110141111112202322232020
Tethydraco_regalis ????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????????014111????????????????
Lonchodraco_giganteus 1??????????????????00??200000-?0??1??0000???????????????????????????010?????110?01?0?????1000101000000001?????????????????????????????????????????????
Ikrandraco_avatar 10100000000110????10001200000-00000------00-11001?01000311?0--0-00?00100101211000112?0-00100010100?0000001111101?????????????2????41??2112201??2??????
Ornithocheirus_simus 10??????????????????0??100?00-?0????????????????????????????????????0?0??????????????0-?0100010000001100??????????????????????????????????????????????
Cimoliopterus_cuvieri 1???????????????????0??100010-?0??1?20000???????????????????????????011??????????????0-?0?000???000???000?????????????????????????????????????????????
Camposipteus_nasutus 1????????????????????0?10001110????????00???????????????????????????011??????????????0-?0?000???000???000?????????????????????????????????????????????
Hamipterus_tianshanensis_ 10100020000?1011111000110001110000121001100-?????001010211?0--0-001001101012010000--10-0011001000000110001111101????1?????00????1041002?12?0??????????
Iberodactylus_andreui ????????????????????0??100?111?0??1?10011???????????????????????????011??????????????????110010000001100??????????????????????????????????????????????
Tropeognathus_mesembrinus 101000200001101011100?110001110000102000000-?????0100102110100100010021010110100010100-011?00100000011010?1111011????11???002???????10211?????????????
AMNH_22555 10101020000110101110011100012?0000102000000-1100101011021101001000100110101????001?100-0111101000000110101111101?1??21112100????1041102112??1?????????
Anhanguera_blittersdorffi 1010102000011010111001110001210000102000000-?????01011021101001000100110101?????0101?0-011110100000011010?????????????????????????????????????????????
Anhanguera_piscator 1010102000011?1????001110001210000102000000-1100101011021101001000100?1010120100010100-0111101000000110101111101?1012111210022111041102112??1?????2120
Liaoningopterus_gui 101?0?2000011???????0??1000121?00?1020000??????????0110211??????????0110????0?0??10100-011110100000011010111?1????????????????????????????????????????
Siroccopteryx_moroccensis 1???????????????????0??0000113?0??1??0000???????????????????????????011??????????????0-???1001000000??010?????????????????????????????????????????????
Uktenadactylus_wadleighi 1???????????????????0??0000113?0?????0000???????????????????????????011??????????????0-???100?0?000???010?????????????????????????????????????????????
Coloborhynchus_clavirostris 1???????????????????0??0000113?0?????0000???????????????????????????011??????????????0-???100???000???010?????????????????????????????????????????????
Ludodactylus_sibbicki 10100000000110?????001110001??0000???000000-1100101001021101161100?00110??1201000101?0-011110100000011110?????????????????????????????????????????????
Guidraco_venator 10?0000000011??????001110001??0000???000000-?????0?0010211?1161100?00?10?0?201000?--?0-011110100000011110?1111????????????????????????????????????????
Caulkicephalus_trimicrodon 10????0??0??????????01?100?12?00??1020000??????????????????11611????011??0???????????0-0??110????00????1??????????????????????????????????????????????
Cearadactylus_atrox 10100??0010?101????0?11100?121?00?1020000?????00??1011?211??????????011010?2010000--00-01111010000001101??????????????????????????????????????????????
Maaradactylus_kellneri 101?0?2001011???????0??1000?21?00?1020000?0?1??????0110211?1????????0110??????0??10100-01111010000001101??????????????????????????????????????????????
Linlongopterus_jennyae 101000?0000?1??????0001200000-00000------00-???????11?0311????????????0????1?11000--?20001000101011000000?????????????????????????????????????????????
Mimodactylus_libanensis_gen._et_sp._nov. ?????????????030??000??200000-?0??????????0-????????????????????00??01001011111000--?20001000111011000000???????0?00111?1?002211??4???211?2013222221??
Haopterus_gracilis 1010???????????????0001200000-?0000------00-?????0????????????????????0???12111?00--020001000111011000000???11??0???111?1?0?22???04???21?????3?22???20
Hongshanopterus_lacustris ?????????????03001000?1200??0-?0???????--0???????????????1??????00??0100101??????????200000001100120000001?11101??????????????????????????????????????
Liaoxipterus_brachyognathus ????????????????????0??1??????????????????????????????????????????????0????1?10000--?21?00000110012000000?????????????????????????????????????????????
Istiodactylus_latidens 1011000111111??????0000100010-00000------00-11100011211330?0--0-001?0?00?011110000--?21000000110012100000?111?011???111020002?1?10411?2112????????????
Istiodactylus_sinensis 1011000111111??????0000100010-00000------00-?????011211330?0--0-00?00300???111?000--?210000001100121000001??11??1???1?1???0??21???4???211???13122??1??
Nurhachius_ignaciobritoi 10110020?10?1??????0001200010-00000------00-??1??01111033000--??????0300?0?2111000--?21000000110012000000??111011???1?10200?2211?041??2112??1312??21??
Nurhachius_luei 1011002001?11??????0001200010-00000------00-0----0????031?00----?0?????????2111000--?20000000110012000000?1?110???????????????????????????????????????
Luchibang_wuke 10?????????????????00??0000?0-?00?0------????????????????????????????????????1100????21000000110012000000?????????????????????????????????????????????
Luchibang_postcrania ??????????????????????????????????????????????????????????????????????????????????????????????????????????1?11?????????02010?201??5[1 2]??1??????312232120
;
cnames
{0 External_naris_(or_nasoantorbital_fenestra),_position_relative_to_the_premaxilla_ main_part_dorsal_to_the_ventral_margin_of_the_premaxilla main_part_displaced_posterior_to_the_premaxilla;
{1 External_naris,_dorsoventrally_compressed absent present;
{2 External_naris_and_antorbital_fenestra,_configuration_ separated confluent,_forming_a_nasoantorbital_fenestra;
{3 External_naris_and_antorbital_fenestra_(or_nasoantorbital_fenestra),_ventral_margin_length_relati ve_the_skull_length shorter_than_40%_of_the_skull_length_ longer_than_40%_of_the_skull_length;
{4 Antorbital_(or_nasoantorbital)_fenestra,_posterior_margin,_shape_ straight_ concave_;
{5 Nasoantorbital_(or_antorbital)_fenestra_extending_dorsal_to_the_orbit_ absent present;
{6 Orbit,_shape_ subcircular quadrangular_(broad_base) piriform_(dorsoventrally_elongated)_;
{7 Orbit,_comparatively_small absent_ present_;
{8 Ventral_margin_of_the_orbit closed open;
{9 Orbit,_position middle_of_the_skull,_with_the_ventral_margin_of_the_orbit_below_the_middle_of_the_antorbital_(or_nasoantorbital)_fenestra_and_the_dorsal_margin_of_the_orbit_above_the_dorsal_margin_of_the_antorbital_(or_nasoantorbital)_fenestra high_in_the_skull,_with_the_ventral_margin_of_the_orbit_the_same_level_or_above_the_middle_ of_the_antorbital_(or_nasoantorbital)_fenestra low_in_the_skull,_with_the_entire_orbit_lower_than_the_dorsal_margin_of_the_antorbital_(or_nasoantorbital)_fenestra;
{10 Suborbital_opening absent_ present_;
{11 Lower_temporal_fenestra,_shape_ comparatively_broad,_with_extensive_subhorizontal_ventral_margin piriform,_with_dorsal_portion_wider_than_ventral__ reduced_(slit-like);
{12 Lower_temporal_fenestra,_position_relative_to_orbit posterior_to_orbit reaches_under_posterior_margin_of_orbit;
{13 Choanae,_separation separated_by_vomer confluent;
{14 Postpalatine_fenestra,_shape_ quadrangular/subtriangular oval egg-shaped elongated_eggshaped_ kite-shaped,_rounded_margin elliptical reduced,_slit-like;
{15 Secondary_subtemporal_fenestra absent present;
{16 Interpterygoid_fenestra,_size_ORDERED extremely_reduced smaller_than_subtemporal_fenestra larger_than_subtemporal_fenestra;
{17 Interpterygoid_fenestra,_shape_ compressed_laterally broad,_longer_than_wide compressed_anteroposteriorly,_wider_than_long round;
{18 Pterygoid_fenestra absent_ present;
{19 Upper_and_lower_jaw,_marked_gap_during_occlusion absent_ present;
{20 Upper_and_lower_jaw,_shape laterally_compressed_ comparatively_broad_;
{21 Skull,_main_part_of_dorsal_margin,_curvature_excluding_cranial_crest nearly_straight concave_ convex;
{22 Length_of_the_rostrum_(pm-naof)_relative_to_the_skull_length_(pm-sq)_ORDERED reduced_ elongated_(about_or_less_than_half_of_skull_length)_ extremely_elongated_(more_than_half_of_skull_length);
{23 Rostrum,_anterior_end,_shape_ flat_surface rounded pointed sharp_tip;
{24 Rostral_end_of_premaxillae/maxillae_downturned absent_ present_;
{25 Rostrum,_distinct_concavity_on_occlusal_surface absent present;
{26 Rostrum,_anterior_portion_forming_a_high_ossified_plate absent present;
{27 Rostrum,_anterior_tip_with_a_slight_dorsal_reflection absent present;
{28 Premaxilla,_anterior_expansion absent present;
{29 Premaxilla,_anterior_expansion,_shape_in_horizantal_plane_ absent elliptical anteriorly_expanded quadrangular;
{30 Premaxilla,_posterior_dorsal_process,_curved_upward absent present;
{31 Premaxillae,_anterior_end_rodlike absent present;
{32 Premaxillary_process_separating_the_external_nares,_thickness_ wide narrow_;
{33 Premaxilla,_posterodorsal_margin_of_nasoantorbital_fenestra_(including_nasal),_width_ wide thin;
{34 Premaxillary_sagittal_crest absent present;
{35 Premaxillary_sagittal_crest_position confined_to_the_anterior_portion_of_the_skull_ starting_anterior_to_the_anterior_margin_of_the_nasoantorbital_fenestra,_extending_beyond_occipital_region starting_at_about_the_anterior_margin_of_the_nasoantorbital_fenestra,_reaching_the_skull_roof_above_the_orbit_but_not_extending_over_the_occipital_region starting_close_or_at_the_anterior_portion_of_the_skull_and_extended_over_the_occipital_region starting_at_the_posterior_half_of_the_nasoantorbital_fenestra starting_at_the_middle_part_of_the_nasoantorbital_fenestra_and_extended_over_the_occipital_region_;
{36 Premaxillary_sagittal_crest,_shape striated,_low_with_a_nearly_straight_dorsal_margin striated,_high_with_a_nearly_straight_dorsal_margin round_dorsal_margin,_bladeshaped smooth,_expanded_anteriorly_and_forming_a_low_rod-like_extension_posteriorly smooth,_starting_low_anteriorly_and_very_expanded_posteriorly striated,_low,_convex_dorsal_margin;
{37 Premaxillary_crest,_elongated_dorsal_premaxillary_spike-like_projection_ absent_ present_;
{38 Premaxillary_crest,_distinct_expansion_on_the_anterior_part_ absent present;
{39 Premaxillary_crest,_concentric_striae_on_the_anterior_region absent present;
{40 Premaxillary_crest,_anterior_expansion_of_the_anterior_margin absent present;
{41 Maxilla,_posterior_ventral_expansion_ absent_ present_;
{42 Maxilla-nasal_contact absent present;
{43 Maxilla-nasal_contact,_broadness broad narrow;
{44 Nasal_descending_process absent present;
{45 Nasal_descending_process,_position_ placed_laterally placed_medially absent;
{46 Nasal_descending_process,_length_ long_(almost_reaching_the__ventral_margin_of_the_skull)_ORDERED short knob-like_(extremely_reduced) absent;
{47 Nasal_descending_process,_orientation_ inclined_anteriorly subvertical;
{48 Nasal_descending_process,_lateral_foramen_ absent_ present_;
{49 Lacrimal,_extensive_fenestration absent_ present_;
{50 Lacrimal,_orbital_process_ absent_ present_;
{51 Jugal,_lacrimal_process_base,_width_ broad narrow;
{52 Jugal,_lacrimal_process,_inclination_ORDERED inclined_anteriorly subvertical inclined_posteriorly;
{53 Jugal,_presence_of_pronounced_ridge_on_the_lateral_side absent present;
{54 Jugal,_posterior_process,_orbital_process absent present;
{55 Quadrate,_inclination_relative_to_ventral_margin_of_skull_ORDERED anteriorly subvertical inclined_about_120°_posteriorlyned_about_120°_backwards_ inclined_about_150°_posteriorly;
{56 Cranio-mandibular_articulation,_position_relative_to_orbit_ORDERED _posterior_to_posterior_margin_of_orbit under_center_of_orbit_ under_anterior_margin_of_the_orbit anterior_to_anterior_margin_of_orbit;
{57 Helical_jaw_joint absent_ present_;
{58 Frontal,_anterior_portion_rugose absent present;
{59 Frontal,_ossified_crest absent present;
{60 Frontal,_ossified_crest,_position_ORDERED confined_to_posterior_end_of_skull starting_above_orbit starting_on__posterior_half_of_nasoantorbital_fenestra;
{61 Frontal,_ossified_crest,_shape_ low,_blunt short._spike-like,_dorsally_deflected spikelike,_directed_posteriorly narrow,_broad,_directed_posteriorly low,_broad_base,_fans-shaped high,_broad_base,_crown-shaped high,_broad_base,_casqued-shaped high,_broad,_directed_posteriorly,_at_least_doubling_shight_of_skull_above_orbit absent;
{62 Parietal,_ossified_crest absent present;
{63 Parietal,_ossified_crest,_shape_ blunt_ constituting_the_base_of_the_posterior_portion_of_the_cranial_crest expanded,_with_rounded_margin;
{64 Posterior_region_of_the_skull_rounded_with_the_squamosal_displaced_ventrally absent_ present_;
{65 Supraoccipital does_not_extend_backwards_ extends_backwards_;
{66 Supraoccipital,_foramen_ absent_ present_;
{67 Paroccipital_processes,_expanded_distal_ends_ absent_ present_;
{68 Foraminae_piercing_the_anterior_portion_of_the_palate,_numerous absent present;
{69 Palatal_occlusal_surface_ORDERED smooth discrete_palatal_ridge,_tapering_anteriorly_ strong_palatal_ridge,_tapering_anteriorly strong_palatal_ridge,_confined_to_the_posterior_portion_of_the_palate;
{70 Palate,_dorsal_deflection absent present;
{71 Palate,_slight_expansion_close_to_the_anterior_margin_of_the_nasoantorbital_(or_naris_+_antorbital)_fenestra absent present;
{72 Maxilla_and_internal_naris,_contact_ absent_ present_;
{73 Palatines,_shape_ broad thin_bars;
{74 Basisphenoid_body,_length_ shorter_than_wide longer_than_wide;
{75 Mandibular_rostral_end,_extension_of_the_contact_surface_of_opposing_dentaries_ORDERED short,_limited_to_the_tip short,_extended_posteriorly_less_than_30%_of_mandible_length long,_up_to_55%_the_mandible_length long,_extended_over_55%_of_mandible_length;
{76 Mandibular_rostral_end,_odontoid_process absent present;
{77 Mandibular_rostral_end,_opposing_dentaries_ unfused fused;
{78 Mandibular_rostral_end,_shape_ rounded pointed sharp_tip;
{79 Dentary,_dorsal_margin,_distinct_posterior_eminence_close_to_the_separation_of_mandibular_rami absent present;
{80 Tip_of_the_dentary_projected_anteriorly absent_ present_;
{81 Dentary_ossified_sagittal_crest absent present;
{82 Dentary_ossified_sagittal_crest,_position_ confined_to_the_anterior_third_of_the_lower_jaw extinding_close_to_the_middle_portion_of_the_lower_jaw;
{83 Dentary_ossified_sagittal_crest,_shape_ shallow blade-like deep,_broad_in_lateral_view elongated_ridge;
{84 Dentary,_posteroventral_fossa absent present;
{85 Teeth,_position_and_presence present,_evenly_distributed_along_the_jaws_ teeth_absent_from_the_anterior_portion_of_the_jaws_ confined_to_the_anterior_part_of_the_jaws jaw_toothless;
{86 Teeth,_confined_to_the_anterior_part_of_the_jaws confined_to_the_anterior_half_of_the_jaws confined_to_the_anterior_quarter_of_the_jaws;
{87 Maxillary_teeth,_largest_positioned_posteriorly_ absent_ present_;
{88 Teeth,_shape_variation isodont heterodont;
{89 Teeth,_anterior,_height_versus_width_proportion more_than_twice_their_width less_than_twice_their_width;
{90 Teeth,_anterior,_marked_variation_in_size absent present;
{91 Teeth,_upper_jaw,_variation_in_the_size_of_the_anterior_teeth_with_the_5th_and_6th_smaller_than_the_4th absent_ present_;
{92 Teeth,_base_broad_and_oval absent_ present_;
{93 Teeth,_serrated present absent;
{94 Teeth,_cingulum absent present;
{95 Teeth,_peg-like_(cone-shaped) absent_ present;
{96 Teeth,_small_needle-shaped absent present;
{97 Teeth,_labiolingually_compressed_crowns absent present;
{98 Teeth,_labiolingually_compressed_crowns,_compression_ORDERED not_compressed slightly_compressed strongly_compressed;
{99 Teeth,_sharp_carinae absent present;
{100 Teeth,_elongated absent present;
{101 Teeth,_striated absent present;
{102 Teeth,_curvature_of_the_toothline absent present;
{103 Teeth,_first_pair_above_second_pair absent present;
{104 Alveoli,_lateral_platform absent present;
{105 Atlas_and_axis unfused_ fused_;
{106 Cervical_vertebrae,_postexapophyses_ absent_ present_;
{107 Mid-cervical_vertebrae,_centrum,_lateral_foramen_ absent_ present_;
{108 Mid-cervical_vertebrae,_length_ORDERED short,_sub-equal_in_length_ longer_than_wide,_with_length_less_than_3_times_width elongated,_with_length_more_than_3_times_width extremely_elongate;
{109 Mid-cervical_vertebrae,_ribs present_ absent_;
{110 Mid-cervical_vertebrae,_neural_spines,_height_ORDERED tall low extremely_reduced_or_absent;
{111 Mid-cervical_vertebrae,_neural_spines,_shape_ blade-shaped spike-shaped_ ridge;
{112 Notarium absent_ present_;
{113 Caudal_vertebrae,_quantity_ more_than_15_ 15_or_less_;
{114 Caudal_vertebrae,_zygapophyses_forming_rod-like_ossified_processes_ absent present;
{115 Proximal_caudal_vertebrae_centrum,_centrum_shape_ single douplex;
{116 Scapula,_length_relative_to_coracoid_length_ORDERED subequal_or_longer_than_coracoid_ scapula_shorter_than_coracoid_(1_>_sca/cor_>_0.80)_ substantially_shorter_than_coracoid_(sca/cor_<_0.80);
{117 Scapula,_proximal_end_ elongated_ sub-oval_;
{118 Scapula,_shape_ elongated_ stout,_with_constructed_shaft_;
{119 Coracoid,_proximal_end,_shape flattened oval;
{120 Coracoid,_sternal_articulation_ORDERED no_developed_articulation articulation_surface_straight_or_slightly_concave articulation_surface_strongly_concave;
{121 Coracoid,_sternal_articulation,_posterior_expansion_ absent present;
{122 Coracoid,_ventral_margin,_deep_flange absent present;
{123 Coracoid,_broad_tubercle_on_ventroposterior_margin absent present;
{124 Cristospine,_shape_ absent_ shallow_and_elongated_ deep_and_short;
{125 Humerus,_proportional_length_relative_to_the_metacarpal_IV_(hu/mcIV)_ORDERED hu/mcIV_>_2.50_ 1.50_<_hu/mcIV_<_2.50_ 0.40_<_hu/mcIV_<_1.50 hu/mcIV_<_0.40;
{126 Humerus,_proportional_length_relative_to_the_femur_(hu/fe)_ORDERED hu/fe_<0.80_ 1.4_>_hu/fe>___0.80_ hu/fe_>_1.40;
{127 Humerus_plus_ulna,_proportional_lengths_relative_to_the_femur_plus_tibia_(hu+ul/fe+ti)_ORDERED humerus_plus_ulna_about_0.80%_or_less_of_femur_plus_tibia_length_(hu+ul/fe+ti_<_0.80)_ humerus_plus_ulna_larger_than_0.80%_of_femur_plus_tibia_length_(hu+ul/fe+ti_>_0.80)_;
{128 Humerus,_proximal_end,_small_foramen_on_dorsal_surface_distal_to_proximal_articulation absent present;
{129 Humerus,_proximal_end,_foramen_on_ventral_surface_close_to_proximal_margin_ absent_ present_;
{130 Humerus,_deltopectoral_crest,_shape_ reduced,_positioned_close_to_the_humerus_shaft enlarged,_proximally_placed,_with_almost_straight_proximal_margin_ enlarged,_hatchet_shaped,_proximally_placed enlarged,_hatched_shaped,_positioned_further_down_the_humerus_shaft enlarged,_warped long,_proximally_placed,_curving_ventrally;
{131 Humerus,_medial_(=_ulnar)_crest_ reduced_ directed_posteriorly_ massive,_with_a_developed_proximal_ridge;
{132 Humerus,_distal_articulation,_shape_ oval_or_D-shaped_ subtriangular_;
{133 Humerus,_between_distal_condyles,_pneumatic_foramen absent present;
{134 Ulna_and_radius,_diameter_at_midshaft_ORDERED subequal_ diameter_of_radius_about_half_that_of_ulna diameter_of_radius_less_than_half_that_of_ulna;
{135 Proximal_syncarpal,_large_posterodistal_process absent present;
{136 Proximal_syncarpal,_shape_(proximal_view) quadrangular_or_irregular pentagonal;
{137 Distal_syncarpals,_shape_(distal_view)_ irregular form_rectangular_unit_ form_triangular_unit;
{138 Pteroid_ORDERED absent_ shorter_than_half_the_length_of_the_ulna_ longer_that_half_the_length_of_the_ulna;
{139 Pteroid,_proximal_articulation,_expanded_in_right_angle_with_shaft absent present;
{140 Metacarpals_I_-_III,_relation_with_carpus_ORDERED articulating_with_carpus_ metacarpal_I_articulates_with_carpus,_metacarpals_II_and_III_reduced_ not_articulating_with_carpus;
{141 Manual_digit_IV_first_phalanx,_proportional_length_relative_to_metacarpal_IV_(ph1d4/mcIV)_ORDERED both_small_and_reduced_ ph1d4/mcIV>4.0 4.0>ph1d4/mcIV>2.0 '2.0>_ph1d4/mcIV>1.0' 'ph1d4/mcIV<_1.0';
{142 Manual_digit_IV_first_phalanx,_proportional_length_relative_to_tibiotarsus_(ph1d4/ti)_ORDERED ph1d4_reduced_ ph1d4_elongated_and_less_than_twice_the_length_of_ti_(ph1d4/ti_smaller_than_2.00) ph1d4_elongated_about_or_longer_than_twice_the_length_of_ti_(ph1d4/ti_subequal/larger_than_2.00);
{143 Manual_digit_IV_second_phalanx,_proportional_length_relative_to_first_phalanx_(ph2d4/ph1d4)__ORDERED both_short_or_absent_ elongated_with_second_phalanx_about_the_same_size_or_longer_than_first_(ph2d4/ph1d4_larger _than_1.00)_ elongated_with_second_phalanx_up_to_30%_shorter_than_first_(ph2d4/ph1d4_between_0.70_-_1.00) elongated_with_second_phalanx_more_than_30%_shorter_than_first_(ph2d4/ph1d4_smaller_than_0.70);
{144 Manual_digit_IV_third_phalanx,_proportional_length_relative_to_first_phalanx_(ph3d4/ph1d4)__ORDERED both_short_or_absent_ ph3d4_about_the_same_length_or_larger_than_ph1d4_ ph3d4_shorter_than_ph1d4;
{145 Proportional_length_of_the_forth_phalanx_of_manual_digit_IV_relative_to_the_first_phalanx_of_manual_digit_IV_(ph4d4/ph1d4)_ORDERED both_short_or_absent both_elongated,_with_the_forth_phalanx_longer_than_the_first_(ph4/d4>1.00) both_elongated,_with_the_forth_phalanx_the_same_length_or_shorter,_but_longer_than_35%_the_length_of_the_first both_elongated,_with_the_forth_phalanx_less_than_35%_the_length_of_the_first;
{146 Femur,_length_relative_to_metacarpal_IV_length_(fe/mcIV)__ORDERED femur_at_least_twice_the_metacarpal_IV_length_(fe_?_mcIV_>_2.00) femur_longer_but_less_than_twice_the_length_of_metacarpal_IV_(1.00_<_fe/mcIV_<_2.00)_ femur_about_the_same_length_or_shorter_than_metacarpal_IV_(fe/mcIV_<_1.00);
{147 Metatarsal_III,_proportional_length_relative_to_tibia_length_ more_than_30%_of_tibia_length_ less_than_30%_of_tibia_length_;
{148 Pedal_digit_V,_number_of_phalanges__ORDERED with_four_phalanges_ with_2_phalanges_ with_1_or_no_phalanx_(extremely_reduced);
{149 Pes,_second_phalanx_of_digit_V,_shape__ reduced_or_absent_ elongated,_straight_ elongated,_curved elongated,_very_curved_(boomerang_shape);
;
ccode + 16 22 46 52 55 56 60 75 98 108 110 116 120 125 126 127 134 140 141 142 143 144 145 146 148*;
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comments 0
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REFERENCES
Goloboff, P.A., Farris, J.S., and Nixon, K.C., 2008, TNT, a free program for phylogenetic analysis: Cladistics, v. 24, p.774-786.
https://doi.org/10.1111/j.1096-0031.2008.00217.x
Hone, D.W., Fitch, A.J., Ma, F. and Xu, X., 2020. An unusual new genus of istiodactylid pterosaur from China based on a near complete specimen. Palaeontologia Electronica, 23(1), pp.1-43.
https://doi.org/10.26879/1015
Kellner, A.W., Caldwell, M.W., Holgado, B., Vecchia, F.M.D., Nohra, R., Sayão, J.M. and Currie, P.J., 2019. First complete pterosaur from the Afro-Arabian continent: insight into pterodactyloid diversity. Scientific Reports, 9(1), pp.1-9.
https://doi.org/10.1038/s41598-019-54042-z
