SUPPLEMENTAL DATA
S1.1. Changes to the Kellner et al. (2019) Matrix:
The matrix of Kellner et al. (2019) was the basis of the phylogenetic analysis of this study, but a number of changes were made to the matrix used here. All provided numbers corresponding to character numberings in the new matrix are provided in italics and within parentheses.
Some characters in the original matrix were pseudoreplicates and so removed. Characters 134 was a pseudoreplicate of characters 125 (125), 126 (126), and 127 (127), and Character 146 was a pseudoreplicate of characters 144 (143) and 145 (144).
A number of characters should be considered ordered because they transition through multiple states (e.g. short > medium > large). However, these characters were left unordered in the original matrix, and so they were specified as ordered in the altered matrix used here. These characters were as follows: their 16, 22, 46, 52, 55, 56, 60, 75, 98, 108, 110, 116, 120, 125, 126, 127, 134, 135, 142, 143, 144, 145, 146, 147, 148, and 150 (16, 22, 46, 52, 55, 56, 60, 75, 98, 108, 110, 116, 120, 125, 126, 127, 134, 140, 141, 142, 143, 144, 145, 146, and 148, respectively).
S1.2. TNT Data Matrix for Total (Cranial & Postcranial) Phylogenetic Analysis of Hone et al., (2020):
xread
150 63
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 101????????????????000?0000?0-?00?0------00-???????????????????????????????1?1100????21000000110012000000?1?11????????102000?201??40??1??????212222120
;
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*;
proc /;
comments 0
;
S1.3. TNT Data Matrix for Postcranial Characters-Only Phylogenetic Analysis of Hone et al., (2020)
xread
45 44
Ornithosuchus_woodwardi 000000000000000000000000000000000000000000000
Herrerasaurus_ischigualastensis 000000000000000000000000000000000000000000020
Scleromochlus_taylori ???00??0000??0?000???00??0???0???00?0000000??
Anurognathus_ammoni ???0???01000?0???0??0110?100000??1001122?0011
Rhamphorhynchus_muensteri 00000000010000010001111002000100?100122221012
Darwinopterus_linglongtaensis ???10??001?0?00?000?1110?100?10??100311111022
Darwinopterus_robustodens ???1???001?0?00??0??111??100?10??10031111102?
Pterodactylus_antiquus 0002110010000001000121100500?100?100312222020
Dsungaripterus_weii 1111100110?000???0?2210??5?001001??13122?2120
Noripterus_parvus ?11??0??????????????????????0?001????????????
Keresdrakon_vilsoni ?111?00?????00020011?0?01520?1???????????????
Tupuxuara_leonardii 11111001???00002001?2100152011001?01313??2???
Caupedactylus_ybaka ???????????00002001????015200????????????????
Caiuajara_dobruski ?1111000?0?00002?01?21?11520?1??120??????????
Tapejara_wellnhoferi ?1111000?0?00002001?210115200100120?31???2120
Sinopterus_dongi ?1?11000???0000200??211?152??100?20?3122321??
Huaxiapterus_corollatus ?1?11??0???000???0??211??5??????????3132321??
Huaxiapterus_benxiensis ?1?11??0????????????2????????????20?3122?2120
Eopteranodon_lii ????????????????????21???5??????????3?22?2???
Chaoyangopterus_zhangi ?102110????000???10?200??????100?20?3132?21?0
Shenzhoupterus_chaoyangensis ?1021??????0?0???1??200??5???100????3122321??
Jidapterus_edentus_ ?1?2110??????0???1??200??5?0?100????312232120
Azhdarcho_lancicollis 11031221???????????????015?0?????????????????
Quetzalcoatlus_sp. 11031221?0?00002010?20?0152001001???4132?2?20
Zhejiangopterus_linhaiensis 1??31221?0?000???10?200??5???100?20?413??2???
Nyctosaurus_gracilis 11?110111000?00100013110130?011122123222?20??
Muzquizopteryx_coahuilensis ???11011????1????????11??30??111?21???????1??
Pteranodon_longiceps 111110111011101200012110141111112202322232020
Tethydraco_regalis ???????????????????????014111????????????????
Ikrandraco_avatar 1111101?????????????2????41??2112201??2??????
Hamipterus_tianshanensis_ 1111101????1?????00????1041002?12?0??????????
Tropeognathus_mesembrinus ?1111011????11???002???????10211?????????????
AMNH_22555 1111101?1??21112100????1041102112??1?????????
Anhanguera_piscator 1111101?1012111210022111041102112??1?????2120
Liaoningopterus_gui 111?1????????????????????????????????????????
Guidraco_venator ?1111????????????????????????????????????????
Mimodactylus_libanensis_gen._et_sp._nov. ???????0?00111?1?002211??4???211?2013222221??
Haopterus_gracilis ???11??0???111?1?0?22???04???21?????3?22???20
Hongshanopterus_lacustris 1?11101??????????????????????????????????????
Istiodactylus_latidens ?111?011???111020002?1?10411?2112????????????
Istiodactylus_sinensis 1??11??1???1?1???0??21???4???211???13122??1??
Nurhachius_ignaciobritoi ??111011???1?10200?2211?041??2112??1312??21??
Nurhachius_luei ?1?110???????????????????????????????????????
Luchibang_wuke ?1?11????????102000?201??40??1??????212222120
;
cnames
{0 Atlas_and_axis unfused_ fused_;
{1 Cervical_vertebrae,_postexapophyses_ absent_ present_;
{2 Mid-cervical_vertebrae,_centrum,_lateral_foramen_ absent_ present_;
{3 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;
{4 Mid-cervical_vertebrae,_ribs present_ absent_;
{5 Mid-cervical_vertebrae,_neural_spines,_height_ORDERED tall low extremely_reduced_or_absent;
{6 Mid-cervical_vertebrae,_neural_spines,_shape_ blade-shaped spike-shaped_ ridge;
{7 Notarium absent_ present_;
{8 Caudal_vertebrae,_quantity_ more_than_15_ 15_or_less_;
{9 Caudal_vertebrae,_zygapophyses_forming_rod-like_ossified_processes_ absent present;
{10 Proximal_caudal_vertebrae_centrum,_centrum_shape_ single douplex;
{11 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);
{12 Scapula,_proximal_end_ elongated_ sub-oval_;
{13 Scapula,_shape_ elongated_ stout,_with_constructed_shaft_;
{14 Coracoid,_proximal_end,_shape flattened oval;
{15 Coracoid,_sternal_articulation_ORDERED no_developed_articulation articulation_surface_straight_or_slightly_concave articulation_surface_strongly_concave;
{16 Coracoid,_sternal_articulation,_posterior_expansion_ absent present;
{17 Coracoid,_ventral_margin,_deep_flange absent present;
{18 Coracoid,_broad_tubercle_on_ventroposterior_margin absent present;
{19 Cristospine,_shape_ absent_ shallow_and_elongated_ deep_and_short;
{20 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;
{21 Humerus,_proportional_length_relative_to_the_femur_(hu/fe)_ORDERED hu/fe_<0.80_ 1.4_>_hu/fe>___0.80_ hu/fe_>_1.40;
{22 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)_;
{23 Humerus,_proximal_end,_small_foramen_on_dorsal_surface_distal_to_proximal_articulation absent present;
{24 Humerus,_proximal_end,_foramen_on_ventral_surface_close_to_proximal_margin_ absent_ present_;
{25 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;
{26 Humerus,_medial_(=_ulnar)_crest_ reduced_ directed_posteriorly_ massive,_with_a_developed_proximal_ridge;
{27 Humerus,_distal_articulation,_shape_ oval_or_D-shaped_ subtriangular_;
{28 Humerus,_between_distal_condyles,_pneumatic_foramen absent present;
{29 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;
{30 Proximal_syncarpal,_large_posterodistal_process absent present;
{31 Proximal_syncarpal,_shape_(proximal_view) quadrangular_or_irregular pentagonal;
{32 Distal_syncarpals,_shape_(distal_view)_ irregular form_rectangular_unit_ form_triangular_unit;
{33 Pteroid_ absent_ shorter_than_half_the_length_of_the_ulna_ longer_that_half_the_length_of_the_ulna;
{34 Pteroid,_proximal_articulation,_expanded_in_right_angle_with_shaft absent present;
{35 Metacarpals_I_-_III,_relation_with_carpus_ORDERED articulating_with_carpus_ metacarpal_I_articulates_with_carpus,_metacarpals_II_and_III_reduced_ not_articulating_with_carpus;
{36 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';
{37 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);
{38 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);
{39 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;
{40 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;
{41 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);
{42 Metatarsal_III,_proportional_length_relative_to_tibia_length_ more_than_30%_of_tibia_length_ less_than_30%_of_tibia_length_;
{43 Pedal_digit_V,_number_of_phalanges__ORDERED with_four_phalanges_ with_2_phalanges_ with_1_or_no_phalanx_(extremely_reduced);
{44 Pes,_second_phalanx_of_digit_V,_shape_ reduced_or_absent_ elongated,_straight_ elongated,_curved elongated,_very_curved_(boomerang_shape);
;
ccode + 3 5 11 15 20 21 29 35 36 37 38 39 40 41 43 *;
proc /;
comments 0
;
S2. Consistency of the specimen
The specimen is unusual for an istiodactylid pterosaur in showing postcranial features more consistent with members of other clades of pterodactyloid pterosaur that are common in the Jehol beds of China. Given that previous specimens are known to have been modified by fossil dealers it has been suggested that this specimen may represent a chimera with an istiodactylid head placed onto an azhdarchid postcranium. Evidence for this hypothesis is assessed below.
In favour of the idea that ELDM 1000 may be a chimera is that the fact that the colour / texture of the cranial elements are not entirely consistent with that of the postcranium, and the unusual distribution of characters between the head and body. With regards to the first point, such changes are also noted in other Jehol-type specimens with some parts often being preserved rather differently to others. For example the head of a specimen of Microraptor (IVPP 17972A) is rather different in colour and preservation to the rest of the specimen and parts of Longchengpterus are much darker than others. While this may in part be due to the manner of preparation of the material, it does not mean these are chimeras. Similarly, although such character changes are discrepancies between two anatomical modules of an animal are very rare, at least one pterosaur group, the Wukongopteridae show just such a distribution of features. Certainly both lines of evidence are important, but neither is definitive.
A number of pieces of data support the idea that the specimen is genuine. The skull of the animal is approximately the correct size for an istiodactylid pterosaur being c. 15% of the wingspan of the animal (total length, allowing of the missing portion) compared to c. 18% for Istiodactylus (based on figure 15.5 Witton, 2013, p.149) and also appears to belong to a juvenile animal given the disarticulation of the posterior part of the cranium. Finding a consistently sized and aged cranium (and indeed is at least very similar if not identical to the postcranium in colour and texture) from a second specimen would be very difficult given the variation seen in preservation of Jehol fossils, supporting the idea that the skull does belong with the postcranium.
If the skull is from a separate animal, then either the bones of this have been incised into a large slab of matrix containing the postcranium, or matrix containing the skull has been added to the main piece. In either case, the spaces between the skull and postcranium are the critical zones and if there have been alterations to the material it should show here. The matrix as a whole is variable, as is common for such specimens, especially when this size, and thus is consider equivocal in this context. However, despite the variation and inevitable crack and breaks, the surface as a whole is generally consistent with itself, and certainly there is nothing distinctive about the region of matrix that include the cranium. Some cracks and breaks have had minor repairs but there is no indication of major repairs or addition of glues, consolidants or other artificial work to any part of the specimen.
Several simple tests were carried out on the material to check for any evidence that the cranium, or the matrix containing this part, were added to the rest of the material. First, the edge of the maxilla where it terminates off the end of the slab was examined visually, and then a small amount of preparation work was carried out at this location. Neither the examination or removal of matrix suggested that there were any breaks or cracks in the matrix that might indicate that the maxilla had been cut into existing matrix.
There is some small amount of glue on the tip of the maxilla (apparently to consolidate the broken tip) but removal of some of this showed that it did not extend far into the matrix, and there were no issues detected outside of the glued area. Additionally, visual examination of the edge of the specimen around the whole upper left quadrant showed no unusual patterns or breaks or inconsistencies between layers of the matrix that would indicate modification to the matrix. The slab is only around 10 mm thick and as a result somewhat fragile, however the edges could be shifted to expose the underside of the main slab and this was also examined around the edge of the upper left quadrant, and again, no issues were detected. There is a pair of large cracks in the specimen though the skull and the cervicals, and also a number of holes across the specimen. The latter are consistent with some surface cracking and damage to the matrix (this is common in Jehol material) and these appear across the specimen, even areas considered quite genuine (e.g. near the right wrist) and thus should not impinge on the validity of the cranium where some similar holes exist, though not directly next to the bone where this might be expected if there has been alterations to the specimen. In the cases of the cracks, no inconsisencies can be found at their margins or along the edges and both are similar to one another in how they affect the matrix and bones. The conclusion here is that the maxilla is naturally within this part of the matrix and the bones have not been cut into an existing piece of matrix.
Similarly, examination and preparation in three distinct locations (between the scapula and mandible, between the right ramus of the mandible and 6th cervical, and finally in the space between the left ramus of the mandible, maxilla and seventh cervical vertebra) by DWEH and an independent observer revealed no evidence of tampering. There was no undercutting of the matrix, no glue or consolidants, and the matrix was entirely consistent in these areas and reached the bone naturally. Between the mandible, maxilla and 7th cervical, the matrix was removed and revealed bone-to-bone contact between each element.
Preparation of these areas and that at the tip of the maxilla was then compared to other parts of the matrix. Several points across the matrix were chosen and subjected to simple scrapes similar to those carried out above to dig into the matrix below the surface layer. These areas were chosen as locations very unlikely to have been altered during any work to add the skull to the postcranium and thus most likely to be correct representatives of the matrix. Again, no inconsistencies were found. Preparation of one of the areas that did have glue on it, showed this to be very difference in appearance to the nature of the matrix prepared in key areas noted above.
Finally, there was a careful visual check (with the naked eye, and both 10X and 30X lenses) of the major elements of the skull and postcranium in places where they touch show that they are continuous. That is, the bones meld into one another where they meet. There are continuous parts of the mandible with the right scapula, with the mandible and maxilla and a cervical (as noted above), and finally with the maxilla and a cervical. This is similar, if not identical to, the joins also seen between multiple cervicals and other postcranial elements. Therefore the mandible, maxilla and postcranium are continuous with one another in what is effectively therefore an unbroken series of elements. There are also some minor cracks on the specimen that appear to run through the bones and matrix quite naturally and link together disparate parts of the specimen. If the skull and / or surrounding matrix had been added to this thin slab, there should be a break somewhere between the two parts and any force sufficient to crack the material should be directed here, and not pass across the break and continue with a natural and normal break across numerous elements, again suggesting these are genuinely and normally connected.
Collectively, these provide very strong evidence that the specimen is genuine and unaltered. Although the skull is an imperfect match, it is largely consistent with the postcranium in size, ontogenetic status, colour and texture. There is no indication that the maxilla (and by extension the skull) has been incised into the matrix, nor any indication that one piece of matrix has been added to another. Areas where modifications to either the bones or matrix would be expected if the head did not match the rest of the specimen also show no indications of changes and the matrix appears to be entirely natural around the bones in numerous parts of the specimen and the surface, edge and underside of the slab appeared consistent. Most critically, as far as can be determined there is a continuum of the bony parts of skull and postcranium with bones meeting and even melding together. This would seem difficult if not absolutely impossible to do convincingly. Preparation in key areas that might expose work done and carefully hidden did not reveal inconsistencies or errors, but instead additional connections between elements.
We cannot absolutely confirm that the specimen is genuine since it was not collected or prepared under our supervision and understandably suspicion will linger, since there are no absolute tests to determine is a given specimen, or part of it, is genuine. All of the evidence must be assessed and an appropriate conclusion reached, and while additional work (e.g. UV light work, further preparation) would be welcomed, we feel that the data is consistent and the specimen should provisionally be considered genuine. However, although some parts of the anatomical disjunct are unexpected and strong, we have been unable to detect any evidence of tampering on any part of the specimen, and indeed have found great consistency especially with the contact between elements.
REFERENCE
Witton MP, 2013. Pterosaurs: natural history, evolution, anatomy. Princeton University Press.
S3. Humeral and Hindlimb Length Data
Length data (in mm) was retained from Elgin (2014). All Pterodactyloidea data was used with the exception of several indeterminate taxa and excluding some of the smallest specimens of archaeopterodactyloids that represent small juveniles. The measurements for Nurhachius ignaciobritoi were added, taken from Wang et al., (2005).
| Clade/Grade | hu | HL | |
| Luchibang wuke | Istiodactylid | 110 | 355 |
| Nurhachius ignaciobritoi | Istiodactylid | 121 | 262 |
| Boreopterus cuiae | Pteranodontoidea | 79 | 164 |
| Arthurdactylus conandoylei | Pteranodontoidea | 230 | 424 |
| Coloborhynchus robustus | Pteranodontoidea | 290 | 615 |
| Coloborhynchus spielbergi | Pteranodontoidea | 290 | 629.6 |
| Pteranodon sp. | Pteranodontidae | 134 | 354 |
| Pteranodon sp. | Pteranodontidae | 247 | 565 |
| Pteranodon sp. | Pteranodontidae | 191 | 446.5 |
| Pteranodon sp. | Pteranodontidae | 168 | 416 |
| Pteranodon sp. | Pteranodontidae | 171 | 385.5 |
| Muzquizopteryx coahuilensis | Pteranodontidae | 80.5 | 187 |
| Nyctosaurus ‘bonneri’ | Pteranodontidae | 97 | 204 |
| Nyctosaurus gracilis | Pteranodontidae | 87 | 214 |
| Nyctosaurus gracilis | Pteranodontidae | 87 | 214 |
| Nyctosaurus sp. | Pteranodontidae | 72.3 | 155.5 |
| Cycnorhamphus suevicus | Archaeopterodactyloidea | 65.5 | 199 |
| Pterodactylus antiquus | Archaeopterodactyloidea | 31.5 | 83 |
| Pterodactylus antiquus | Archaeopterodactyloidea | 43.7 | 104 |
| Pterodactylus antiquus | Archaeopterodactyloidea | 31.7 | 76 |
| Pterodactylus antiquus | Archaeopterodactyloidea | 32.5 | 77 |
| Pterodactylus antiquus | Archaeopterodactyloidea | 41 | 98.5 |
| Beipiaopterus chenianus | Archaeopterodactyloidea | 68 | 144 |
| Ctenochasma gracilis | Archaeopterodactyloidea | 38.5 | 90 |
| Ctenochasma gracilis | Archaeopterodactyloidea | 17 | 35.2 |
| Ctenochasma gracilis | Archaeopterodactyloidea | 16.9 | 36.5 |
| Ctenochasma gracilis | Archaeopterodactyloidea | 19.2 | 37 |
| Ctenochasma gracilis | Archaeopterodactyloidea | 19.5 | 38.3 |
| Gnathosaurus subulatus | Archaeopterodactyloidea | 23 | 60 |
| Gnathosaurus subulatus | Archaeopterodactyloidea | 23 | 57 |
| Gnathosaurus subulatus | Archaeopterodactyloidea | 25 | 69.5 |
| Gnathosaurus subulatus | Archaeopterodactyloidea | 26 | 74 |
| Gnathosaurus subulatus | Archaeopterodactyloidea | 28 | 82 |
| "Pterodactylus" longicollum | Archaeopterodactyloidea | 78 | 248 |
| Pterodaustro guinazui | Archaeopterodactyloidea | 80 | 150 |
| Germanodactylus cristatus | Archaeopterodactyloidea | 56 | 56.5 |
| Germanodactylus rhamphastinus | Archaeopterodactyloidea | 49.5 | 130 |
| Germanodactylus rhamphastinus | Archaeopterodactyloidea | 60 | 159 |
| Huaxiapterus corollatus | Azhdarchoidea | 79.7 | 247.5 |
| Huaxiapterus jii | Azhdarchoidea | 79 | 241 |
| Sinopterus dongi | Azhdarchoidea | 58.5 | 178 |
| Sinopterus gui | Azhdarchoidea | 35.5 | 96.1 |
| Chaoyangopterus zhangi | Azhdarchoidea | 93 | 338 |
| Eoazhdarcho liaoxiensis | Azhdarchoidea | 90 | 254 |
| Shenzhoupterus chaoyangensis | Azhdarchoidea | 66 | 241 |
| Zhejiangopterus linhaiensis | Azhdarchoidea | 137 | 487 |
REFERENCES
Elgin R. A. 2014. Palaeobiology, Morphology, and flight characteristics of pterodactyloid pterosaurs. PhD Thesis, University of Heidelberg. 273pp.
Wang, X.-L., Kellner, A.W.A., Zhou, Z.-H., Campos, D.A., 2005. Pterosaur diversity and faunal turnover in Cretaceous terrestrial ecosystems in China. Nature 437, 875-879.
S4. Supporting figures
FIGURE S1. Close up of the premaxillary teeth of ELDM 1000 in lateral view. Scale bar has 1 cm divisions.
FIGURE S2. Close up of the mandibular teeth and symphysis of ELDM 1000 in dorsal view. Scale bar has 1 cm divisions.
FIGURE S3. Close up of the fish preserved under the dorsal ribs of ELDM 1000. Scale bar has 1 cm divisions.
FIGURE S4. Close up of part of the large patch of probable skin on ELDM 1000. Scale bar has 1 cm divisions.
