#NEXUS [!This data set was downloaded from TreeBASE, a relational database of phylogenetic knowledge. TreeBASE has been supported by the NSF, Harvard University, Yale University, SDSC and UC Davis. Please do not remove this acknowledgment from the Nexus file. Generated on November 14, 2025; 3:44 GMT TreeBASE (cc) 1994-2008 Study reference: Whiting M., Carpenter J., Wheeler Q., & Wheeler W. 1997. The strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology. Systematic Biology, 46(1): 1-68. TreeBASE Study URI: http://purl.org/phylo/treebase/phylows/study/TB2:S382] [ The following blocks are input data for analysis step 433 ] BEGIN TAXA; TITLE TaxaForAnalysisStep433; DIMENSIONS NTAX=111; TAXLABELS Acyrthosiphon_pisum Agrion_maculatum Agulla_sp. Anisomorpha_buprestoides Apoica_sp. Archaeognatha Ascalapha_odorata Bareogonalos_canadensis Bittacus_chlorostigmus Bittacus_strigosus Blaberus_sp. Blattodea Boreus_coloradensis Buenoa_sp. Caenocholax_fenyesii Caenochrysis_doriae Camponotus_ligniperda Cerastipsocus_venosus Chalepoxenus_muellerianus Chrysoperla_plorabunda Coleoptera Collembola Colpocaccus_posticatus Corydalus_cognatus Crawfordia_n._sp. Ctenocephalides_canis Cultus_decisus Cybister_fimbriolatus Dasymutilla_gloriosa Dennyus_hirudensis Dermaptera Diptera Doronomyrmex_kutteri Drosophila_melanogaster Elenchus_japonica Embioptera Ephemerella_sp. Ephemerida Epyris_sepulchralis Evania_appendigaster Galleria_mellonella Harpagoxenus_sublaevis Hartigia_cressonii Hemerobius_stigmata Hemiptera Hemitaxonus_sp. Hydropsyche_sparna Hymenoptera Hypogastrura_sp. Hystrichopsylla_schefferi Ichneumon_sp. Labidura_riparia Laphria_sp. Lepidoptera Leptothorax_acervorum Libellula_pulchella Lolomyia_texana Lygus_lineolaris Mantis_religiosa Mantispa_pulchella Mantodea Mecoptera Megaloptera Megarcys_stigmata Melanoplus_sp. Meloe_proscarabaeus Merope_tuber Mesopolobus_sp. Monobia_quadridens Myrmeleon_immaculatus Myrmeleon_sp. Mythicomyia_atra Neuroptera Octinodes_sp. Odonata Oecetis_avara Oligotoma_saundersii Ophion_sp. Orchopeas_leucopus Orthoptera Orussus_thoracicus Panorpa_latipennis Papilio_troilus Peridista_linea Phasmida Photuris_pennsylvanicus Phthiraptera Plecoptera Polistes_dominulus Polistes_fuscatus Priacma_serrata Priocnemus_oregana Psocoptera Pycnopsyche_lepida Raphidioptera Rhipiphorus_fasciatus Saldula_pallipes Siphonaptera Strepsiptera Taeniothrips_inconsequens Temnothorax_recedens Tenebrio_molitor Tetraopes_tetropthalmus Thysanoptera Tipula_sp. Trichoptera Trigoniopthalmus_alternatus Triozocera_mexicana Xenos_pecki Xenos_vesparum Xyloryctes_faunus ; END; BEGIN CHARACTERS; [! TreeBASE Matrix URI: http://purl.org/phylo/treebase/phylows/matrix/TB2:M2818] TITLE Appendix_3; LINK TAXA = TaxaForAnalysisStep433; DIMENSIONS NCHAR=176; FORMAT DATATYPE=Standard SYMBOLS= "0 1 2 3 4" MISSING=? GAP= -; CHARSTATELABELS 1 Cerci / Absent 'present, originating from appen' simplified unsegmented, 2 Malphigian_tubules / 'Developed, many (>6)' lost four, 3 Anterior_tentorium / Present reduced, 4 Posterior_tentorium / Absent bearing_arms arms_fused, 5 Ovipositor / Absent present vestigial modified fused, 6 Caudal_filament / Absent 'present, long' short, 7 Mandibular_articulation / Monocondylic dicondylic anterior_articulation_fixed, 8 Gonangulum_in_ovipositor_base / Absent present, 9 Origin_of_ventral_mandibular_an / On_endoskeletal_plates on_tentorium, 10 Fulturae / Present absent, 11 Postoccipital_ridge / Absent internal_ridge_lateral_only continuous, 12 Tracheal_commisures_and_connect / Not_developed_in_abdomen developed_in_abdomen, 13 Amniotic_cavity / Absent Open Closed, 14 'Median fusion (at least basal p' / Absent present, 15 Paired_female_genital_openings / Absent present, 16 Two_pairs_of_wings / Absent present absent_in_adult_females, 17 Two_coxal_proprioreceptor_organ / Absent present, 18 Basal_wing_brace / Absent present, 19 Media_stem / Absent present, 20 Superlinguae / Absent present, 21 Habitat_of_nymphs / Terrestrial aquatic, 22 Subimago / Present absent, 23 Tracheation / Anterior_only arch, 24 Posterior_tracheation_of_leg / Absent present, 25 Direct_spiracular_musculature / Absent present, 26 'Tentorio-lacinial muscle' / Present absent, 27 'Tentorio-mandibular muscles' / Several_bundles one, 28 Loss_of_pterothoracic_muscles / No yes, 29 Sperm_transfer / Indirect copulation 'indirect, using claspers', 30 Wing_flexion / Absent present, 31 Third_valvulae_forming_sheath / Absent present, 32 Anal_furrow / Absent present, 33 Enlarged_hindwing_vannus / Small enlarged, 34 Jugal_bar / Absent present, 35 Ocelli / Present absent_in_immatures absent_in_adults median_ocellus_absent median_ocellus_absent_in_adults, 36 Metameric_testes_ducts / Present absent, 37 Male_gonocoxopidites_IX / Articulated not_articulated, 38 Metaspina / Present absent, 39 Phallomeres / Normal reduced, 40 Male_styli / Expressed supressed, 41 Segmental_arteries / Absent present, 42 Female_broodcare_behavior / Absent present, 43 Trochantin / Absent present 'trochantin-episternal sulcus pr', 44 Sclerotization_of_forewing / Membranous sclerotized, 45 Vannus_pleated / No yes, 46 Two_cervical_sclerites / No yes, 47 Prognathy / Absent present gula_present, 48 Dorsal_paraglossa_flexor / Normal, 49 Phallic_rudiments_produce_gonop / No yes, 50 Prominent_precostal_field / Absent present, 51 Discoid_pronotum / Absent present enlarged, 52 Pleural_sutures / Not_slanting_forward slanting_forward, 53 Indirect_wing_muscles / Developed reduced, 54 Metathoracic_tergosternal_wing / Present weak_or_suppressed, 55 'Coxa conical, backwards-directe' / No yes, 56 Phallomeres / Symmetrical asymmetrical, 57 Shape_of_mesotrochantin / No yes, 58 1st_branch_of_Cu / Not_grooved Grooved, 59 Separate_coxopleuron / Yes no, 60 Several_male_accessory_glands_a / No yes, 61 Median_ventral_excurrent_ostia / Absent present, 62 Mesothoracic_basisternal_fold / Absent present, 63 Premental_lobes / Free fused, 64 Tarsi / Not_subdivided '5-segmented' '3-segmented', 65 Sternum_I / Developed Reduced absent, 66 'Dorsal-flexor of paraglossae' / Absent present, 67 'Second profurca-spinasternal mu' / Absent present, 68 Female_sternum_VII / Small large forming_vestibulum, 69 Sex_determination / 'XX-XY' 'XX-XO', 70 Corporotentorium / Absent present perforation_through_which_circu, 71 Anterior_teeth_of_proventriculu / Absent present, 72 Proventriculus_teeth_with_secon / Absent present, 73 Tarsal_plantulae / Absent present, 74 Ootheca / Absent present ootheca_hardened_after_depositi, 75 First_axillary_sclerite_attache / Absent present, 76 Sperm_axoneme_sheaths / Absent present two_dense_sheaths_surround_the, 77 Abdominal_ganglia / Unconcentrated one, 78 Lacinia / Broad slender_rod stylets, 79 Flagella_of_spermatozoa / Single two, 80 Gonangulum_fused_with_tergum_IX / No yes, 81 Cibarium / Absent present, 82 Ovarioles / Telotrophic_or_panoistic polytrophic, 83 Basal_part_of_antennal_flagello / Absent present, 84 Mandibles / Broad stylettiform, 85 Sclerotized_rings_between_anten / Absent present, 86 Metamorphosis / Incomplete complete, 87 Larval_eyes / Carried_over_to_adult not_carried_over_to_adult, 88 Wing_rudiments / Evaginated_prior_to_penultimate 'evaginated at larval-pupal molt', 89 Appearance_of_external_genitali / Prior_to_penultimate_molt penultimate_molt, 90 Cruciate_cervical_muscles / Present absent, 91 Female_genitalia / Gonopohyses_8_developed_Gonopohyses_9_discrete Reduced_fused_not_discrete, 92 Pretarsal_claw_of_larval_leg / Paired unpaired, 93 Silk_secretion_from_larval_labi / Absent present, 94 Eruciform_larvae / Absent present, 95 'Outer tergo-coxal remotor muscl' / Present absent, 96 Telomere / Present absent, 97 Abdominal_limb_buds / Repressed, 98 Gastric_caeca / Present lost, 99 Meron / Absent present, 100 Cryptosterny / Absent present, 101 Flight_by_posteromotorism / No yes, 102 'Intrinsic musculature in (fused' / Absent present, 103 Connection_of_metepimeron_to_an / Weakly_or_not_connected firmly_connected, 104 Reinforcement_of_tergum_II_acro / Absent present, 105 Pleural_muscle_inserted_on_firs / No yes, 106 Larval_stipes / Entire transversely_divided, 107 'Larval cranial cardo-promotor,' / Present absent, 108 Cranial_antagonist_of_the_primi / Absent present, 109 Female_heterogamety / Absent present, 110 'Double Y-shaped fusion of anal' / Absent present, 111 Achiasmatic_oogenesis / Absent present, 112 Preholocentric_chromosomes / Absent present, 113 Pterothoracic_furcal_arms_fused / Absent present, 114 Pair_of_glands_opening_on_stern / Absent present, 115 Outer_accessory_filaments_of_sp / Thin 'very stout (ca. 300 Å diam.)', 116 Vestiture_on_wing_surfaces_betw / Absent dense_vestiture_of_long_setae scales, 117 'Dagger-like mandible with anter' / Absent present articulation_lost, 118 'Prelabium without endite lobes-' / No yes, 119 Labial_palp_segments / Five two absent, 120 Tentorial_muscles_of_prelabium / Present lost, 121 Structure_of_posterior_notal_wi / Not_modified modified, 122 'Pleural ridge-scutum muscle ins' / No yes, 123 Transverse_muscle_between_profu / Absent present, 124 Lateral_labral_retractor / Present absent, 125 Tentorial_adductors_of_cardo_in / Present absent, 126 Hypopharyngeal_muscles_in_larva / Present retractor_of_hypopharynx_and_ve 'loss of mouth-angle retractors', 127 Larvae / Podous apodous, 128 Extrinsic_labral_muscles / Present absent, 129 Proventriculus_with_specialized / Absent present, 130 Flagellum_of_spermatozoon_coile / Absent present, 131 Procoxa_with_condylus_fitting_p / No yes, 132 'Coxa-body articulation' / None 'pleural, mobile' 'pleural, fixed' tricondylic, 133 Larval_labium / Not_prehensile prehensile, 134 Posterior_tracheal_larval_gills / Absent present, 135 Lateral_cervical_sclerite_in_th / No yes, 136 Pteropleura_tilted_backward_wit / No yes, 137 Male_accessory_copulatory_organ / Absent present, 138 Tufted_larval_tracheal_gills / Absent 'present, lateral', 139 Eversible_vesicle_on_abdominal / Absent present, 140 Transverse_stipital_muscle / Present absent, 141 Vertebrate_ectoparasites / No yes, 142 Labium_ensheathing_mandibular_a / No yes, 143 Maxillary_palpi_lost / No yes, 144 Maxillary_and_mandibular_stylet / Absent present, 145 'Larval piercing-sucking tubes' / Absent present, 146 Elytra / Absent present, 147 Abdominal_segments_VIII_and_IX / No yes, 148 Abdominal_sterna_I_and_II_membr / No yes, 149 Mesothorax_and_metathorax_fused / No yes, 150 Insect_endoparasitism / Absent present, 151 Forewing / Absent present, 152 Antennae_flabellate / No yes, 153 'Male eyes protruding, with omma' / No yes, 154 Hamuli / Absent present, 155 Notocoxal_muscles / Present absent, 156 Volsella / Absent present, 157 Venom_production_by_female_acce / Absent present, 158 Male_antennae_serving_as_access / No yes, 159 Ctenidium_on_prothorax / Absent present, 160 Salivary_channels_in_lacinial_s / No yes, 161 Body_laterally_compressed / No yes, 162 Clypeus_and_labrum_fused / No yes, 163 Stylar_organ_of_the_male_claspe / Absent present, 164 'Male abdominal segment IX ring-' / No yes, 165 Spermathecal_opening_separate_f / No yes, 166 Halteres / Absent present, 167 Hypopharynx_styletlike / No yes, 168 Labial_palpi_forming_labellum / No yes, 169 Larval_spiracles_without_closin / No yes, 170 'Larvae apneustic, respiration e' / No yes, 171 Anal_prolegs / Absent present, 172 Larval_antennae_papillae / No yes, 173 Adult_labium_and_hypopharynx_fu / No yes, 174 Abdominal_tergum_X_bilobed / No yes, 175 Corporotentorium_with_median_po / No yes, 176 Ventral_nerve_cord_solid / No yes, ; MATRIX [ 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 ] [ . . . . . . . . . . . . . . . . . ] Archaeognatha 10021100001011000??000?000000?0???000000000000000?00??000?000000000000000000000000000000000000000000?00000000?000000000000000000000100000010000000000000000000000000000000000000 Blattodea 10021021112121011000011111111111103111001011110000211111001101011002121111000000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Coleoptera 10021021112121011000011111111111011111000010000010000000111000010000010000000001000001111110000100011000000000000000000000000000000300000001000001111000000000000000000000000000 Collembola 0101000?000000000??100?000000?0???000000000000000?00??000?000000000000000000000000000000000000000000?00000000?000000000000000000000000000010000000000000000000000000000000000000 Dermaptera 100220201121200110000111111111?1102111010121111000111100001?01020001010010010000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Diptera 1002002?11212101100001111111111???1111000010?00010000000111000010000010000000200000001111001111001110000111100000000211111111110000300000001000000000000000000000001011110000000 Embioptera 2002002?1121210210100111111111?1002111110120012100010000000110120110010010100000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Ephemerida 1002011?11212011111110000000100000000000000??00000000000000000010000010000000000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Hemiptera 02021021112121011000011111111111011111000010000010000000101000022000010000001111000110000000000000000000000000000000002000000000000200000001011100000000000000000000000?00000000 Hymenoptera 10021021112121011000011111111111011111000010000010000000111000010000010000000000000001111001110011010000000000000000000000000000000300000001000000000000011110000000000000000000 Lepidoptera 1002002?112121011000011111111111014111000010000010000000111000010000010000000000000001111001111011110000111111111112000000000000000300000001000000000000000000000000000000000111 Mantodea 10021021112121011000011111111111100111001011110000011111001101011002121112000000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Mecoptera 1002002?112121011000011111111111011111000010000010000000111000010000010000000000000001111001111011110000111100000000111111111201111300000001000000000000000000000111100000000000 Megaloptera 10024021112121011000111111111111011111000010000010000000111000010000010000000001000001111110000011110111010000000000000000000000000300000001000000000000000000000000000000000000 Neuroptera 10024021112121011000011111111111011111000010000010000000111000010000010000000001000001111110000001110100010000000000000000000000000300000001000010000000000000000000000000000000 Odonata 1002122111212101111011111111200000000000000??00000000000000000020000010000000000000000000000000000000000000000000000000000000000000211111001000000000000000000000000000000000000 Orthoptera 20023021112121011010011111111111100111000011111001000000001100110000010010000000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Phasmida 30023021112121011010011111111111100111000011111101000000001100110110010010110000000000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Phthiraptera 02021021112121001??001?111111?1???111100001??0001?00??001?100002200001000002111111100000000000000000?00000000?000000000000000000000200000001100000000000000000000000000000000000 Plecoptera 1012002?1121200110101111111111?1100011110020010000000000110010120000010010000000000000000000000000000000000000000000000000000000000200000100000000000000000000000000000000000000 Psocoptera 02021021112121011000011110111111011111000010000010000000101000022000010000001111111000000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Raphidioptera 10024021112121011000011111111111011111000010000010000000111000010000010000000001000001111110000001110111000000000000000000000000000300000001000000000000000000000000000000000000 Siphonaptera 1002002?112121001??001?111111?1???111100001??0001?00??001?100001000001000000020000000111100111100111?000?1010?000000?001??111111111300000001100000000000000001111000000000000000 Strepsiptera 1010002??1212102100001111??111?1011111000000000010000000?11000010000010000000?000000010010??001?0?1110000???00000000112?00????0??00300000001000000000111100000000001000?00000000 Thysanoptera 02021021112121011000011111111111011111000010000010000000101000022000010000001111000110000000000000000000000000000000000000000000000200000001000000000000000000000000000000000000 Trichoptera 1002002?1121210110001111111111110111110000100000100000001110000100000100000000000000011110011110111100001111111111110000000000000003000000010000000000000000000000000000?1111000 ; END; [ The following blocks are output data for analysis step 433 ] BEGIN TREES; TITLE Tb5213; LINK TAXA = TaxaForAnalysisStep433; TREE Fig._5 = [&R] (Collembola,(Archaeognatha,(Ephemerida,(Odonata,(((Plecoptera,Embioptera),((Phasmida,Orthoptera),(Dermaptera,(Blattodea,Mantodea)))),(((Hemiptera,Thysanoptera),(Psocoptera,Phthiraptera))Paraneoptera,((Coleoptera,(Neuroptera,(Megaloptera,Raphidioptera))),(Hymenoptera,((Lepidoptera,Trichoptera),(Strepsiptera,(Diptera,(Mecoptera,Siphonaptera)))Antliophora)Mecopterida))Holometabola))Neoptera))Insecta)); [! TreeBASE tree URI: http://purl.org/phylo/treebase/phylows/tree/TB2:Tr3310] END;