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Insights into the evolution of the desert stink beetles
- 1. Insights into the evolution of the desert stink beetles: Combining phylogenemics and phenotypes (Coleoptera: Tenebrionidae: Amphidorini) M. Andrew Johnston, Kojun Kanda, Aaron D. Smith @MAndrewJohnston
- 2. Desert Stink Beetles Tribe Amphidorini LeConte, 1862 • “Desert Stink Beetles” or “false wireworms” • 253 valid species/subspecies in North America • >400 available names (described species) Eleodimorpha bolcan Blaisdell, 1909 Eleodes armata LeConte, 1851
- 3. Desert Stink Beetles Steneleodes longicollis LeConte, 1851 ‘Head-standing’ behavior
- 4. Desert Stink Beetles Steneleodes longicollis LeConte, 1851 ‘Head-standing’ behavior Paired defensive glands
- 7. Phylotranscriptomic Analysis Generated 27 partial transcriptomes
- 8. Phylotranscriptomic Analysis Generated 27 partial transcriptomes Assembled 5 ortholog sets Totaling 844 loci
- 9. Phylotranscriptomic Analysis Generated 27 partial transcriptomes Assembled 5 ortholog sets Totaling 844 loci Coalescent species-tree reconstruction with ASTRAL-II
- 10. Species-tree analyses in ASTRAL-II • 4 monophyletic groups found consistently Phylotranscriptomic Analysis
- 11. Species-tree analyses in ASTRAL-II • 4 monophyletic groups found consistently • Relationships between clades less clear ? Phylotranscriptomic Analysis
- 12. Amphidora-clade sister to all others Found in more conservative FO-aa and SISRS-1k datasets Phylotranscriptomic Analysis
- 13. Eleodes-clade sister to all others FO-nt SISRS-loci Combined-nt Phylotranscriptomic Analysis
- 14. Preferred topology? Phylotranscriptomic Analysis • Rapid early diversification events?
- 15. Preferred topology? Phylotranscriptomic Analysis Eleodes Species Non-Eleodes Species 5 1 53 20 38 142 0 0 • Rapid early diversification events? • Eleodes Eschscholtz, 1829 is paraphyletic
- 16. Preferred topology? Phylotranscriptomic Analysis • Rapid early diversification events? • Eleodes Eschscholtz, 1829 is paraphyletic • To avoid splitting, everything would become Embaphion Eleodes Species Non-Eleodes Species 5 1 53 20 38 142 0 0
- 17. Amphidorini as of January 2018 • 6 genera and 16 subgenera
- 18. • 6 genera and 16 subgenera • 253 valid sp/ssp (411 described) Amphidorini as of January 2018
- 19. • 6 genera and 16 subgenera • 253 valid sp/ssp (411 described) • 232 within Eleodes Amphidorini as of January 2018
- 20. Phylogenetic scaffold for generic changes Eleodes (New Sense)
- 21. Revised state of Amphidorini • 16 genera and 9 subgenera Torugena Globostyla Exarenula 3 new genera
- 22. Revised state of Amphidorini • 16 genera and 9 subgenera • 11 genus-group changes
- 23. Revised state of Amphidorini • 16 genera and 9 subgenera • 11 genus-group changes • 113 species-group changes
- 24. Why so many similar species? Sympatric species assemblages from Arizona 5 species from mid-elevation
- 25. Why so many similar species? Sympatric species assemblages from Arizona 5 species from mid-elevation Smith et al. 2014, Fig. 2 Müllerian mimicry complex
- 26. Why so many similar species? Apomorphic lineages – Broad phenotypic variation possible Steneleodes_ponderosus_E22 Melaneleodes_tricostatus_E68 Embaphion_depressum_E73 Eleodes_scapularis_E158 Eleodes_labialis_TB15486 Eleodes__Eleodes__loretensis_TB15982 Eleodes_nr__dentipes_TB15474 Eleodes_ponderosus_TB15078 Eleodes_quadricollis_TB15293 Eleodes_rossi_E35 Eleodes__Steneleodes__sp__TB15477 Melaneleodes_anthracinus_E173 Eleodes_spinipes_ventriculosus_E18 Steneleodes_laevigatus_E174 Promus_subnitens_E10 Lariversius_eureka_TB15302 Litheleodes_extr_TXElPaso_E52 Melaneleodes_carb_chihuahuaensis_E136 Steneleodes_pr_coarctatus_E172 Embaphion_glabrum_E50 Steneleodes_hepburni_E8 Chaseleodes_connatus_E58 Eleodes_pilosus_Horn_1870__Eleodes__TB15449 Eleodes_clavicocnis_E83 Lariversius_tibialis_BigDune_E79 Eleodes_sp_Tlaxcala_E132 Eleodes_opacus_TB17183 Eleodes_acuticaudus_LeConte_TB15444 Eleodes_sp_TB15481 Neobaphion_alleni_KKDNA0426 Eleodes_sp4_TB15082 Promus_calcaratus_E163 Metablapylis_nigrinus_E38 Steneleodes_innocens_E41 Litheleodes_extricatus_AZGraham_E63 Blapylis_tenebrosus_E47 Eleodes_sallaei_var_2_TB15075 Blapylis_tribulus_E61 Litheleodes_extr__NavajoNM_E55 Eleodes_spinipes_spinipes_E155 Promus_goryi_E27 Steneleodes_UnknownA_desierto_liones_E170 Cratidus_osculans_E36 Steneleodes_longicollis_E16 Eleodes_sp3_TB15081 Tricheleodes_hirsutus_E164 Lariversius_tibialis_E70 Lariversius_tibialis2_E57 Eleodes_longipilosus_TB15304 Eleodes_giganteus_TB15475 Embaphion_contusum_E46 Tricheleodes_pilosus_E45 Melaneleodes_carbonarius_E14 Eleodes_sp__nr__pilosus_TB15305 Eleodes_peropacus_TB15476 Eleodes_obs_sulcipennis_E11 Metablapylis_dissimilis_E7 Embaphion_elongatum_TB15300 Eleodes_dentipes_TB15297 Promus_oaxaca_sp_E165 Discogenia_scabriculus_E67 Eleodes__Eleodes__armatus_TB13139 Promus_hogei_E40 Eleodes_impolitus_E53 Eleodes_debilis_TB15710 Heteropromus_veterator_E74 Eleodes__Melaneleodes__debilis_TB13393 Steneleodes_sallaei_E169 Eleodes_sp_Oaxaca_E146 Litheleodes_extr_TXWard_E48 Eleodes_longicollis_TB15295 Promus_knullorum_E24 Embaphion_NMSmall_TB17171 Promus_madrensis_E128 Eleodes_sponsus_E39 Eleodes__Steneleodes__TB15303 Blaps_mucronota_TB15113 Blaps_sp1_TB17179 Eleodes_sp5_TB15083 Blapylis_sp_TulareCo_E142 Eleodes__Pseudeleodes__sp__KKDNA0392 Eleodes_sp7_TB15085 Eleodes_barbatus_TB15992 Metablapylis_delicatus_E141 Neobaphion_papula_E157 Litheleodes_extricatus_AZNav_E62 Eleodes_distinctus_TB15079 Eleodes_nr_distinctus_E144 Tricheleodes_barbatus_E59 Caverneleodes_microps_E76 Embaphion_muricatum_E51 Caverneleodes_Mexico_KKDNA0410_Lib0222 Eleodes_impolitus_TB15074 Eleodes_rileyi_TB15994 Omegeleodes_puebla_sp_E167 Steneleodes_ruidus_E33 Metablapylis_n_sp__teelsmarsh_E175 Steneleodes_impolitus_E166 Eleodes_marginatus_TB15890 Eleodes_suturalis_E19 Eleodes_aristatus_TB13389 Promus_alutaceus_E49 Eleodimorpha_bolcan_KKDNA0407_Lib0223 Lariversius_kelso_TB13435 Trogloderus_NMSanJuan_E91 Eleodes__Eleodes__grandicollis_TB15986 Eleodes_sp2_TB15080 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15447 Ardeleodes_tibialis_E145 Promus_spinolae_E32 Ele_sp81_1_TB13394 Promus_striolatus_E80 Eleodes_sp9_TB15114 Eleodes_grandicollis_TB13402 Eleodes_dissimilis_TB15478 Eleodes_loretensis_TB15674 Embaphion_sp1_Carlsbad_TB15451 Trogloderus_costatus_E60 Eleodes_sp8_TB15077 Eleodes_tenuipes_TB15993 Caverneleodes_wynnei_E26 Trogloderus_Olancha_E81 Eleodes__Promus__insularis_TB15981 Eleodes_mexicanus_TB15483 Melaneleodes_carb_obsoletus_E42 Promus_fusiformis_E44 Eleodes_tibialis_TB15112 Blaps_sp3_TB17181 Blapylis_snowi_E31 Promus_insularis_E54 Steneleodes_Hidalgo_sp_E160 Eleodes_easterlai_KKDNA0425 Eleodes_caudiferus_E29 Eleodes_spinipes_spinipes_TB15076 Ele_sp1_TB13388 Omegeleodes_debilis_E64 Caverneleodes_thomasi_E77 Omegeleodes_oaxaca_sp2_E161 Eleodes_spinipes_macrurus_TB15473 Eleodes_acuticaudus_E34 Eleodes_barbatus_TB15989 Eleodes_unknownB_E171 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15446 Eleodes_sp6_TB15084 Eleodes_hirsutus_TB15479 Neobaphion_planipennis_E43 Trogloderus_costatus_costatu_E176 Blapylis_nigropilosus_E90 Eleodes_armatus_E13 Steneleodes_giganteus_E20 Omegeleodes_oaxaca_sp_E159 Eleodes_acutus_TB17182 Melaneleodes_wenzeli_E15 Melaneleodes_carb_omissus_E12 Eleodes_puebla_sp_E162 Steneleodes_mutilatus_E9 Caverneleodes_n_sp_1_Tonto_E25 Steneleodes_gravidus_E30 Eleodes_esch_eschscholtzi_E23 Steneleodes_distinctus_E154 Eleodes_loretensis_TB15485 Eleodes_gracilis_E28 Eleodes_80_E37 Blaps_sp2_TB17180 Eleodes__Eleodes__loretensis_TB15983 Litheleodes_arcuatus_E140 Eleodes_eschscholtzi_E21 Eleodes_tenuipes_E17 Eleodes__Melaneleodes__debilis_TB15988 Steneleodes_angustus_E168 100 100 9 1 100 100 6 2 7 9 9 1 100 4 7 6 0 3 8 100 3 9 4 6 4 6 9 3 4 2 9 8 100 6 5 4 3 100 8 7 9 0 9 0 8 2 100 9 8 6 7 6 7 7 2 100 9 6 1 9 100 6 3 5 1 1 2 5 2 4 0 100 9 9 100 3 8 3 4 1 3 9 4 100 7 6 9 8 9 8 100 9 7 9 8 100 100 8 8 2 1 2 0 7 6 100 100 4 8 8 4 100 9 8 6 5 3 6 100 5 7 5 5 9 2 9 5 100 100 4 0 9 7 7 8 6 0 3 6 100 9 4 3 9 100 100 9 8 1 8 100 9 9 9 9 4 3 7 8 100 100 9 9 100 2 1 5 3 9 9 100 100 3 4 100 9 9 9 3 6 5 100 4 1 8 2 3 8 5 2 4 3 8 9 100 100 4 2 100 2 3 100 100 100 100 6 6 100 100 9 6 100 9 2 5 3 8 2 5 6 8 2 100 100 100 100 9 0 9 8 100 9 9 100 6 7 1 2 100 7 4 100 9 9 9 4 3 0 9 9 100 9 9 100 5 5 100 9 9 100 100 8 2 100 100 100 100 100 100 100 100
- 27. Why so many similar species? Apomorphic lineages – Broad phenotypic variation possible Steneleodes_ponderosus_E22 Melaneleodes_tricostatus_E68 Embaphion_depressum_E73 Eleodes_scapularis_E158 Eleodes_labialis_TB15486 Eleodes__Eleodes__loretensis_TB15982 Eleodes_nr__dentipes_TB15474 Eleodes_ponderosus_TB15078 Eleodes_quadricollis_TB15293 Eleodes_rossi_E35 Eleodes__Steneleodes__sp__TB15477 Melaneleodes_anthracinus_E173 Eleodes_spinipes_ventriculosus_E18 Steneleodes_laevigatus_E174 Promus_subnitens_E10 Lariversius_eureka_TB15302 Litheleodes_extr_TXElPaso_E52 Melaneleodes_carb_chihuahuaensis_E136 Steneleodes_pr_coarctatus_E172 Embaphion_glabrum_E50 Steneleodes_hepburni_E8 Chaseleodes_connatus_E58 Eleodes_pilosus_Horn_1870__Eleodes__TB15449 Eleodes_clavicocnis_E83 Lariversius_tibialis_BigDune_E79 Eleodes_sp_Tlaxcala_E132 Eleodes_opacus_TB17183 Eleodes_acuticaudus_LeConte_TB15444 Eleodes_sp_TB15481 Neobaphion_alleni_KKDNA0426 Eleodes_sp4_TB15082 Promus_calcaratus_E163 Metablapylis_nigrinus_E38 Steneleodes_innocens_E41 Litheleodes_extricatus_AZGraham_E63 Blapylis_tenebrosus_E47 Eleodes_sallaei_var_2_TB15075 Blapylis_tribulus_E61 Litheleodes_extr__NavajoNM_E55 Eleodes_spinipes_spinipes_E155 Promus_goryi_E27 Steneleodes_UnknownA_desierto_liones_E170 Cratidus_osculans_E36 Steneleodes_longicollis_E16 Eleodes_sp3_TB15081 Tricheleodes_hirsutus_E164 Lariversius_tibialis_E70 Lariversius_tibialis2_E57 Eleodes_longipilosus_TB15304 Eleodes_giganteus_TB15475 Embaphion_contusum_E46 Tricheleodes_pilosus_E45 Melaneleodes_carbonarius_E14 Eleodes_sp__nr__pilosus_TB15305 Eleodes_peropacus_TB15476 Eleodes_obs_sulcipennis_E11 Metablapylis_dissimilis_E7 Embaphion_elongatum_TB15300 Eleodes_dentipes_TB15297 Promus_oaxaca_sp_E165 Discogenia_scabriculus_E67 Eleodes__Eleodes__armatus_TB13139 Promus_hogei_E40 Eleodes_impolitus_E53 Eleodes_debilis_TB15710 Heteropromus_veterator_E74 Eleodes__Melaneleodes__debilis_TB13393 Steneleodes_sallaei_E169 Eleodes_sp_Oaxaca_E146 Litheleodes_extr_TXWard_E48 Eleodes_longicollis_TB15295 Promus_knullorum_E24 Embaphion_NMSmall_TB17171 Promus_madrensis_E128 Eleodes_sponsus_E39 Eleodes__Steneleodes__TB15303 Blaps_mucronota_TB15113 Blaps_sp1_TB17179 Eleodes_sp5_TB15083 Blapylis_sp_TulareCo_E142 Eleodes__Pseudeleodes__sp__KKDNA0392 Eleodes_sp7_TB15085 Eleodes_barbatus_TB15992 Metablapylis_delicatus_E141 Neobaphion_papula_E157 Litheleodes_extricatus_AZNav_E62 Eleodes_distinctus_TB15079 Eleodes_nr_distinctus_E144 Tricheleodes_barbatus_E59 Caverneleodes_microps_E76 Embaphion_muricatum_E51 Caverneleodes_Mexico_KKDNA0410_Lib0222 Eleodes_impolitus_TB15074 Eleodes_rileyi_TB15994 Omegeleodes_puebla_sp_E167 Steneleodes_ruidus_E33 Metablapylis_n_sp__teelsmarsh_E175 Steneleodes_impolitus_E166 Eleodes_marginatus_TB15890 Eleodes_suturalis_E19 Eleodes_aristatus_TB13389 Promus_alutaceus_E49 Eleodimorpha_bolcan_KKDNA0407_Lib0223 Lariversius_kelso_TB13435 Trogloderus_NMSanJuan_E91 Eleodes__Eleodes__grandicollis_TB15986 Eleodes_sp2_TB15080 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15447 Ardeleodes_tibialis_E145 Promus_spinolae_E32 Ele_sp81_1_TB13394 Promus_striolatus_E80 Eleodes_sp9_TB15114 Eleodes_grandicollis_TB13402 Eleodes_dissimilis_TB15478 Eleodes_loretensis_TB15674 Embaphion_sp1_Carlsbad_TB15451 Trogloderus_costatus_E60 Eleodes_sp8_TB15077 Eleodes_tenuipes_TB15993 Caverneleodes_wynnei_E26 Trogloderus_Olancha_E81 Eleodes__Promus__insularis_TB15981 Eleodes_mexicanus_TB15483 Melaneleodes_carb_obsoletus_E42 Promus_fusiformis_E44 Eleodes_tibialis_TB15112 Blaps_sp3_TB17181 Blapylis_snowi_E31 Promus_insularis_E54 Steneleodes_Hidalgo_sp_E160 Eleodes_easterlai_KKDNA0425 Eleodes_caudiferus_E29 Eleodes_spinipes_spinipes_TB15076 Ele_sp1_TB13388 Omegeleodes_debilis_E64 Caverneleodes_thomasi_E77 Omegeleodes_oaxaca_sp2_E161 Eleodes_spinipes_macrurus_TB15473 Eleodes_acuticaudus_E34 Eleodes_barbatus_TB15989 Eleodes_unknownB_E171 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15446 Eleodes_sp6_TB15084 Eleodes_hirsutus_TB15479 Neobaphion_planipennis_E43 Trogloderus_costatus_costatu_E176 Blapylis_nigropilosus_E90 Eleodes_armatus_E13 Steneleodes_giganteus_E20 Omegeleodes_oaxaca_sp_E159 Eleodes_acutus_TB17182 Melaneleodes_wenzeli_E15 Melaneleodes_carb_omissus_E12 Eleodes_puebla_sp_E162 Steneleodes_mutilatus_E9 Caverneleodes_n_sp_1_Tonto_E25 Steneleodes_gravidus_E30 Eleodes_esch_eschscholtzi_E23 Steneleodes_distinctus_E154 Eleodes_loretensis_TB15485 Eleodes_gracilis_E28 Eleodes_80_E37 Blaps_sp2_TB17180 Eleodes__Eleodes__loretensis_TB15983 Litheleodes_arcuatus_E140 Eleodes_eschscholtzi_E21 Eleodes_tenuipes_E17 Eleodes__Melaneleodes__debilis_TB15988 Steneleodes_angustus_E168 100 100 9 1 100 100 6 2 7 9 9 1 100 4 7 6 0 3 8 100 3 9 4 6 4 6 9 3 4 2 9 8 100 6 5 4 3 100 8 7 9 0 9 0 8 2 100 9 8 6 7 6 7 7 2 100 9 6 1 9 100 6 3 5 1 1 2 5 2 4 0 100 9 9 100 3 8 3 4 1 3 9 4 100 7 6 9 8 9 8 100 9 7 9 8 100 100 8 8 2 1 2 0 7 6 100 100 4 8 8 4 100 9 8 6 5 3 6 100 5 7 5 5 9 2 9 5 100 100 4 0 9 7 7 8 6 0 3 6 100 9 4 3 9 100 100 9 8 1 8 100 9 9 9 9 4 3 7 8 100 100 9 9 100 2 1 5 3 9 9 100 100 3 4 100 9 9 9 3 6 5 100 4 1 8 2 3 8 5 2 4 3 8 9 100 100 4 2 100 2 3 100 100 100 100 6 6 100 100 9 6 100 9 2 5 3 8 2 5 6 8 2 100 100 100 100 9 0 9 8 100 9 9 100 6 7 1 2 100 7 4 100 9 9 9 4 3 0 9 9 100 9 9 100 5 5 100 9 9 100 100 8 2 100 100 100 100 100 100 100 100 Trogloderus skillmani n.sp.
- 28. Why so many similar species? Apomorphic lineages – Broad phenotypic variation possible Steneleodes_ponderosus_E22 Melaneleodes_tricostatus_E68 Embaphion_depressum_E73 Eleodes_scapularis_E158 Eleodes_labialis_TB15486 Eleodes__Eleodes__loretensis_TB15982 Eleodes_nr__dentipes_TB15474 Eleodes_ponderosus_TB15078 Eleodes_quadricollis_TB15293 Eleodes_rossi_E35 Eleodes__Steneleodes__sp__TB15477 Melaneleodes_anthracinus_E173 Eleodes_spinipes_ventriculosus_E18 Steneleodes_laevigatus_E174 Promus_subnitens_E10 Lariversius_eureka_TB15302 Litheleodes_extr_TXElPaso_E52 Melaneleodes_carb_chihuahuaensis_E136 Steneleodes_pr_coarctatus_E172 Embaphion_glabrum_E50 Steneleodes_hepburni_E8 Chaseleodes_connatus_E58 Eleodes_pilosus_Horn_1870__Eleodes__TB15449 Eleodes_clavicocnis_E83 Lariversius_tibialis_BigDune_E79 Eleodes_sp_Tlaxcala_E132 Eleodes_opacus_TB17183 Eleodes_acuticaudus_LeConte_TB15444 Eleodes_sp_TB15481 Neobaphion_alleni_KKDNA0426 Eleodes_sp4_TB15082 Promus_calcaratus_E163 Metablapylis_nigrinus_E38 Steneleodes_innocens_E41 Litheleodes_extricatus_AZGraham_E63 Blapylis_tenebrosus_E47 Eleodes_sallaei_var_2_TB15075 Blapylis_tribulus_E61 Litheleodes_extr__NavajoNM_E55 Eleodes_spinipes_spinipes_E155 Promus_goryi_E27 Steneleodes_UnknownA_desierto_liones_E170 Cratidus_osculans_E36 Steneleodes_longicollis_E16 Eleodes_sp3_TB15081 Tricheleodes_hirsutus_E164 Lariversius_tibialis_E70 Lariversius_tibialis2_E57 Eleodes_longipilosus_TB15304 Eleodes_giganteus_TB15475 Embaphion_contusum_E46 Tricheleodes_pilosus_E45 Melaneleodes_carbonarius_E14 Eleodes_sp__nr__pilosus_TB15305 Eleodes_peropacus_TB15476 Eleodes_obs_sulcipennis_E11 Metablapylis_dissimilis_E7 Embaphion_elongatum_TB15300 Eleodes_dentipes_TB15297 Promus_oaxaca_sp_E165 Discogenia_scabriculus_E67 Eleodes__Eleodes__armatus_TB13139 Promus_hogei_E40 Eleodes_impolitus_E53 Eleodes_debilis_TB15710 Heteropromus_veterator_E74 Eleodes__Melaneleodes__debilis_TB13393 Steneleodes_sallaei_E169 Eleodes_sp_Oaxaca_E146 Litheleodes_extr_TXWard_E48 Eleodes_longicollis_TB15295 Promus_knullorum_E24 Embaphion_NMSmall_TB17171 Promus_madrensis_E128 Eleodes_sponsus_E39 Eleodes__Steneleodes__TB15303 Blaps_mucronota_TB15113 Blaps_sp1_TB17179 Eleodes_sp5_TB15083 Blapylis_sp_TulareCo_E142 Eleodes__Pseudeleodes__sp__KKDNA0392 Eleodes_sp7_TB15085 Eleodes_barbatus_TB15992 Metablapylis_delicatus_E141 Neobaphion_papula_E157 Litheleodes_extricatus_AZNav_E62 Eleodes_distinctus_TB15079 Eleodes_nr_distinctus_E144 Tricheleodes_barbatus_E59 Caverneleodes_microps_E76 Embaphion_muricatum_E51 Caverneleodes_Mexico_KKDNA0410_Lib0222 Eleodes_impolitus_TB15074 Eleodes_rileyi_TB15994 Omegeleodes_puebla_sp_E167 Steneleodes_ruidus_E33 Metablapylis_n_sp__teelsmarsh_E175 Steneleodes_impolitus_E166 Eleodes_marginatus_TB15890 Eleodes_suturalis_E19 Eleodes_aristatus_TB13389 Promus_alutaceus_E49 Eleodimorpha_bolcan_KKDNA0407_Lib0223 Lariversius_kelso_TB13435 Trogloderus_NMSanJuan_E91 Eleodes__Eleodes__grandicollis_TB15986 Eleodes_sp2_TB15080 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15447 Ardeleodes_tibialis_E145 Promus_spinolae_E32 Ele_sp81_1_TB13394 Promus_striolatus_E80 Eleodes_sp9_TB15114 Eleodes_grandicollis_TB13402 Eleodes_dissimilis_TB15478 Eleodes_loretensis_TB15674 Embaphion_sp1_Carlsbad_TB15451 Trogloderus_costatus_E60 Eleodes_sp8_TB15077 Eleodes_tenuipes_TB15993 Caverneleodes_wynnei_E26 Trogloderus_Olancha_E81 Eleodes__Promus__insularis_TB15981 Eleodes_mexicanus_TB15483 Melaneleodes_carb_obsoletus_E42 Promus_fusiformis_E44 Eleodes_tibialis_TB15112 Blaps_sp3_TB17181 Blapylis_snowi_E31 Promus_insularis_E54 Steneleodes_Hidalgo_sp_E160 Eleodes_easterlai_KKDNA0425 Eleodes_caudiferus_E29 Eleodes_spinipes_spinipes_TB15076 Ele_sp1_TB13388 Omegeleodes_debilis_E64 Caverneleodes_thomasi_E77 Omegeleodes_oaxaca_sp2_E161 Eleodes_spinipes_macrurus_TB15473 Eleodes_acuticaudus_E34 Eleodes_barbatus_TB15989 Eleodes_unknownB_E171 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15446 Eleodes_sp6_TB15084 Eleodes_hirsutus_TB15479 Neobaphion_planipennis_E43 Trogloderus_costatus_costatu_E176 Blapylis_nigropilosus_E90 Eleodes_armatus_E13 Steneleodes_giganteus_E20 Omegeleodes_oaxaca_sp_E159 Eleodes_acutus_TB17182 Melaneleodes_wenzeli_E15 Melaneleodes_carb_omissus_E12 Eleodes_puebla_sp_E162 Steneleodes_mutilatus_E9 Caverneleodes_n_sp_1_Tonto_E25 Steneleodes_gravidus_E30 Eleodes_esch_eschscholtzi_E23 Steneleodes_distinctus_E154 Eleodes_loretensis_TB15485 Eleodes_gracilis_E28 Eleodes_80_E37 Blaps_sp2_TB17180 Eleodes__Eleodes__loretensis_TB15983 Litheleodes_arcuatus_E140 Eleodes_eschscholtzi_E21 Eleodes_tenuipes_E17 Eleodes__Melaneleodes__debilis_TB15988 Steneleodes_angustus_E168 100 100 9 1 100 100 6 2 7 9 9 1 100 4 7 6 0 3 8 100 3 9 4 6 4 6 9 3 4 2 9 8 100 6 5 4 3 100 8 7 9 0 9 0 8 2 100 9 8 6 7 6 7 7 2 100 9 6 1 9 100 6 3 5 1 1 2 5 2 4 0 100 9 9 100 3 8 3 4 1 3 9 4 100 7 6 9 8 9 8 100 9 7 9 8 100 100 8 8 2 1 2 0 7 6 100 100 4 8 8 4 100 9 8 6 5 3 6 100 5 7 5 5 9 2 9 5 100 100 4 0 9 7 7 8 6 0 3 6 100 9 4 3 9 100 100 9 8 1 8 100 9 9 9 9 4 3 7 8 100 100 9 9 100 2 1 5 3 9 9 100 100 3 4 100 9 9 9 3 6 5 100 4 1 8 2 3 8 5 2 4 3 8 9 100 100 4 2 100 2 3 100 100 100 100 6 6 100 100 9 6 100 9 2 5 3 8 2 5 6 8 2 100 100 100 100 9 0 9 8 100 9 9 100 6 7 1 2 100 7 4 100 9 9 9 4 3 0 9 9 100 9 9 100 5 5 100 9 9 100 100 8 2 100 100 100 100 100 100 100 100 Embaphion muricatum (Say, 1824)
- 29. Why so many similar species? Apomorphic lineages – Broad phenotypic variation possible Steneleodes_ponderosus_E22 Melaneleodes_tricostatus_E68 Embaphion_depressum_E73 Eleodes_scapularis_E158 Eleodes_labialis_TB15486 Eleodes__Eleodes__loretensis_TB15982 Eleodes_nr__dentipes_TB15474 Eleodes_ponderosus_TB15078 Eleodes_quadricollis_TB15293 Eleodes_rossi_E35 Eleodes__Steneleodes__sp__TB15477 Melaneleodes_anthracinus_E173 Eleodes_spinipes_ventriculosus_E18 Steneleodes_laevigatus_E174 Promus_subnitens_E10 Lariversius_eureka_TB15302 Litheleodes_extr_TXElPaso_E52 Melaneleodes_carb_chihuahuaensis_E136 Steneleodes_pr_coarctatus_E172 Embaphion_glabrum_E50 Steneleodes_hepburni_E8 Chaseleodes_connatus_E58 Eleodes_pilosus_Horn_1870__Eleodes__TB15449 Eleodes_clavicocnis_E83 Lariversius_tibialis_BigDune_E79 Eleodes_sp_Tlaxcala_E132 Eleodes_opacus_TB17183 Eleodes_acuticaudus_LeConte_TB15444 Eleodes_sp_TB15481 Neobaphion_alleni_KKDNA0426 Eleodes_sp4_TB15082 Promus_calcaratus_E163 Metablapylis_nigrinus_E38 Steneleodes_innocens_E41 Litheleodes_extricatus_AZGraham_E63 Blapylis_tenebrosus_E47 Eleodes_sallaei_var_2_TB15075 Blapylis_tribulus_E61 Litheleodes_extr__NavajoNM_E55 Eleodes_spinipes_spinipes_E155 Promus_goryi_E27 Steneleodes_UnknownA_desierto_liones_E170 Cratidus_osculans_E36 Steneleodes_longicollis_E16 Eleodes_sp3_TB15081 Tricheleodes_hirsutus_E164 Lariversius_tibialis_E70 Lariversius_tibialis2_E57 Eleodes_longipilosus_TB15304 Eleodes_giganteus_TB15475 Embaphion_contusum_E46 Tricheleodes_pilosus_E45 Melaneleodes_carbonarius_E14 Eleodes_sp__nr__pilosus_TB15305 Eleodes_peropacus_TB15476 Eleodes_obs_sulcipennis_E11 Metablapylis_dissimilis_E7 Embaphion_elongatum_TB15300 Eleodes_dentipes_TB15297 Promus_oaxaca_sp_E165 Discogenia_scabriculus_E67 Eleodes__Eleodes__armatus_TB13139 Promus_hogei_E40 Eleodes_impolitus_E53 Eleodes_debilis_TB15710 Heteropromus_veterator_E74 Eleodes__Melaneleodes__debilis_TB13393 Steneleodes_sallaei_E169 Eleodes_sp_Oaxaca_E146 Litheleodes_extr_TXWard_E48 Eleodes_longicollis_TB15295 Promus_knullorum_E24 Embaphion_NMSmall_TB17171 Promus_madrensis_E128 Eleodes_sponsus_E39 Eleodes__Steneleodes__TB15303 Blaps_mucronota_TB15113 Blaps_sp1_TB17179 Eleodes_sp5_TB15083 Blapylis_sp_TulareCo_E142 Eleodes__Pseudeleodes__sp__KKDNA0392 Eleodes_sp7_TB15085 Eleodes_barbatus_TB15992 Metablapylis_delicatus_E141 Neobaphion_papula_E157 Litheleodes_extricatus_AZNav_E62 Eleodes_distinctus_TB15079 Eleodes_nr_distinctus_E144 Tricheleodes_barbatus_E59 Caverneleodes_microps_E76 Embaphion_muricatum_E51 Caverneleodes_Mexico_KKDNA0410_Lib0222 Eleodes_impolitus_TB15074 Eleodes_rileyi_TB15994 Omegeleodes_puebla_sp_E167 Steneleodes_ruidus_E33 Metablapylis_n_sp__teelsmarsh_E175 Steneleodes_impolitus_E166 Eleodes_marginatus_TB15890 Eleodes_suturalis_E19 Eleodes_aristatus_TB13389 Promus_alutaceus_E49 Eleodimorpha_bolcan_KKDNA0407_Lib0223 Lariversius_kelso_TB13435 Trogloderus_NMSanJuan_E91 Eleodes__Eleodes__grandicollis_TB15986 Eleodes_sp2_TB15080 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15447 Ardeleodes_tibialis_E145 Promus_spinolae_E32 Ele_sp81_1_TB13394 Promus_striolatus_E80 Eleodes_sp9_TB15114 Eleodes_grandicollis_TB13402 Eleodes_dissimilis_TB15478 Eleodes_loretensis_TB15674 Embaphion_sp1_Carlsbad_TB15451 Trogloderus_costatus_E60 Eleodes_sp8_TB15077 Eleodes_tenuipes_TB15993 Caverneleodes_wynnei_E26 Trogloderus_Olancha_E81 Eleodes__Promus__insularis_TB15981 Eleodes_mexicanus_TB15483 Melaneleodes_carb_obsoletus_E42 Promus_fusiformis_E44 Eleodes_tibialis_TB15112 Blaps_sp3_TB17181 Blapylis_snowi_E31 Promus_insularis_E54 Steneleodes_Hidalgo_sp_E160 Eleodes_easterlai_KKDNA0425 Eleodes_caudiferus_E29 Eleodes_spinipes_spinipes_TB15076 Ele_sp1_TB13388 Omegeleodes_debilis_E64 Caverneleodes_thomasi_E77 Omegeleodes_oaxaca_sp2_E161 Eleodes_spinipes_macrurus_TB15473 Eleodes_acuticaudus_E34 Eleodes_barbatus_TB15989 Eleodes_unknownB_E171 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15446 Eleodes_sp6_TB15084 Eleodes_hirsutus_TB15479 Neobaphion_planipennis_E43 Trogloderus_costatus_costatu_E176 Blapylis_nigropilosus_E90 Eleodes_armatus_E13 Steneleodes_giganteus_E20 Omegeleodes_oaxaca_sp_E159 Eleodes_acutus_TB17182 Melaneleodes_wenzeli_E15 Melaneleodes_carb_omissus_E12 Eleodes_puebla_sp_E162 Steneleodes_mutilatus_E9 Caverneleodes_n_sp_1_Tonto_E25 Steneleodes_gravidus_E30 Eleodes_esch_eschscholtzi_E23 Steneleodes_distinctus_E154 Eleodes_loretensis_TB15485 Eleodes_gracilis_E28 Eleodes_80_E37 Blaps_sp2_TB17180 Eleodes__Eleodes__loretensis_TB15983 Litheleodes_arcuatus_E140 Eleodes_eschscholtzi_E21 Eleodes_tenuipes_E17 Eleodes__Melaneleodes__debilis_TB15988 Steneleodes_angustus_E168 100 100 9 1 100 100 6 2 7 9 9 1 100 4 7 6 0 3 8 100 3 9 4 6 4 6 9 3 4 2 9 8 100 6 5 4 3 100 8 7 9 0 9 0 8 2 100 9 8 6 7 6 7 7 2 100 9 6 1 9 100 6 3 5 1 1 2 5 2 4 0 100 9 9 100 3 8 3 4 1 3 9 4 100 7 6 9 8 9 8 100 9 7 9 8 100 100 8 8 2 1 2 0 7 6 100 100 4 8 8 4 100 9 8 6 5 3 6 100 5 7 5 5 9 2 9 5 100 100 4 0 9 7 7 8 6 0 3 6 100 9 4 3 9 100 100 9 8 1 8 100 9 9 9 9 4 3 7 8 100 100 9 9 100 2 1 5 3 9 9 100 100 3 4 100 9 9 9 3 6 5 100 4 1 8 2 3 8 5 2 4 3 8 9 100 100 4 2 100 2 3 100 100 100 100 6 6 100 100 9 6 100 9 2 5 3 8 2 5 6 8 2 100 100 100 100 9 0 9 8 100 9 9 100 6 7 1 2 100 7 4 100 9 9 9 4 3 0 9 9 100 9 9 100 5 5 100 9 9 100 100 8 2 100 100 100 100 100 100 100 100 Amphidora littoralis Eschscholtz, 1829 Alice Abela - Bugguide
- 30. Why so many similar species? Conserved phenotype represented in most species Steneleodes_ponderosus_E22 Melaneleodes_tricostatus_E68 Embaphion_depressum_E73 Eleodes_scapularis_E158 Eleodes_labialis_TB15486 Eleodes__Eleodes__loretensis_TB15982 Eleodes_nr__dentipes_TB15474 Eleodes_ponderosus_TB15078 Eleodes_quadricollis_TB15293 Eleodes_rossi_E35 Eleodes__Steneleodes__sp__TB15477 Melaneleodes_anthracinus_E173 Eleodes_spinipes_ventriculosus_E18 Steneleodes_laevigatus_E174 Promus_subnitens_E10 Lariversius_eureka_TB15302 Litheleodes_extr_TXElPaso_E52 Melaneleodes_carb_chihuahuaensis_E136 Steneleodes_pr_coarctatus_E172 Embaphion_glabrum_E50 Steneleodes_hepburni_E8 Chaseleodes_connatus_E58 Eleodes_pilosus_Horn_1870__Eleodes__TB15449 Eleodes_clavicocnis_E83 Lariversius_tibialis_BigDune_E79 Eleodes_sp_Tlaxcala_E132 Eleodes_opacus_TB17183 Eleodes_acuticaudus_LeConte_TB15444 Eleodes_sp_TB15481 Neobaphion_alleni_KKDNA0426 Eleodes_sp4_TB15082 Promus_calcaratus_E163 Metablapylis_nigrinus_E38 Steneleodes_innocens_E41 Litheleodes_extricatus_AZGraham_E63 Blapylis_tenebrosus_E47 Eleodes_sallaei_var_2_TB15075 Blapylis_tribulus_E61 Litheleodes_extr__NavajoNM_E55 Eleodes_spinipes_spinipes_E155 Promus_goryi_E27 Steneleodes_UnknownA_desierto_liones_E170 Cratidus_osculans_E36 Steneleodes_longicollis_E16 Eleodes_sp3_TB15081 Tricheleodes_hirsutus_E164 Lariversius_tibialis_E70 Lariversius_tibialis2_E57 Eleodes_longipilosus_TB15304 Eleodes_giganteus_TB15475 Embaphion_contusum_E46 Tricheleodes_pilosus_E45 Melaneleodes_carbonarius_E14 Eleodes_sp__nr__pilosus_TB15305 Eleodes_peropacus_TB15476 Eleodes_obs_sulcipennis_E11 Metablapylis_dissimilis_E7 Embaphion_elongatum_TB15300 Eleodes_dentipes_TB15297 Promus_oaxaca_sp_E165 Discogenia_scabriculus_E67 Eleodes__Eleodes__armatus_TB13139 Promus_hogei_E40 Eleodes_impolitus_E53 Eleodes_debilis_TB15710 Heteropromus_veterator_E74 Eleodes__Melaneleodes__debilis_TB13393 Steneleodes_sallaei_E169 Eleodes_sp_Oaxaca_E146 Litheleodes_extr_TXWard_E48 Eleodes_longicollis_TB15295 Promus_knullorum_E24 Embaphion_NMSmall_TB17171 Promus_madrensis_E128 Eleodes_sponsus_E39 Eleodes__Steneleodes__TB15303 Blaps_mucronota_TB15113 Blaps_sp1_TB17179 Eleodes_sp5_TB15083 Blapylis_sp_TulareCo_E142 Eleodes__Pseudeleodes__sp__KKDNA0392 Eleodes_sp7_TB15085 Eleodes_barbatus_TB15992 Metablapylis_delicatus_E141 Neobaphion_papula_E157 Litheleodes_extricatus_AZNav_E62 Eleodes_distinctus_TB15079 Eleodes_nr_distinctus_E144 Tricheleodes_barbatus_E59 Caverneleodes_microps_E76 Embaphion_muricatum_E51 Caverneleodes_Mexico_KKDNA0410_Lib0222 Eleodes_impolitus_TB15074 Eleodes_rileyi_TB15994 Omegeleodes_puebla_sp_E167 Steneleodes_ruidus_E33 Metablapylis_n_sp__teelsmarsh_E175 Steneleodes_impolitus_E166 Eleodes_marginatus_TB15890 Eleodes_suturalis_E19 Eleodes_aristatus_TB13389 Promus_alutaceus_E49 Eleodimorpha_bolcan_KKDNA0407_Lib0223 Lariversius_kelso_TB13435 Trogloderus_NMSanJuan_E91 Eleodes__Eleodes__grandicollis_TB15986 Eleodes_sp2_TB15080 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15447 Ardeleodes_tibialis_E145 Promus_spinolae_E32 Ele_sp81_1_TB13394 Promus_striolatus_E80 Eleodes_sp9_TB15114 Eleodes_grandicollis_TB13402 Eleodes_dissimilis_TB15478 Eleodes_loretensis_TB15674 Embaphion_sp1_Carlsbad_TB15451 Trogloderus_costatus_E60 Eleodes_sp8_TB15077 Eleodes_tenuipes_TB15993 Caverneleodes_wynnei_E26 Trogloderus_Olancha_E81 Eleodes__Promus__insularis_TB15981 Eleodes_mexicanus_TB15483 Melaneleodes_carb_obsoletus_E42 Promus_fusiformis_E44 Eleodes_tibialis_TB15112 Blaps_sp3_TB17181 Blapylis_snowi_E31 Promus_insularis_E54 Steneleodes_Hidalgo_sp_E160 Eleodes_easterlai_KKDNA0425 Eleodes_caudiferus_E29 Eleodes_spinipes_spinipes_TB15076 Ele_sp1_TB13388 Omegeleodes_debilis_E64 Caverneleodes_thomasi_E77 Omegeleodes_oaxaca_sp2_E161 Eleodes_spinipes_macrurus_TB15473 Eleodes_acuticaudus_E34 Eleodes_barbatus_TB15989 Eleodes_unknownB_E171 Eleodes_littoralis_Eschscholtz_1831__Amphidora__TB15446 Eleodes_sp6_TB15084 Eleodes_hirsutus_TB15479 Neobaphion_planipennis_E43 Trogloderus_costatus_costatu_E176 Blapylis_nigropilosus_E90 Eleodes_armatus_E13 Steneleodes_giganteus_E20 Omegeleodes_oaxaca_sp_E159 Eleodes_acutus_TB17182 Melaneleodes_wenzeli_E15 Melaneleodes_carb_omissus_E12 Eleodes_puebla_sp_E162 Steneleodes_mutilatus_E9 Caverneleodes_n_sp_1_Tonto_E25 Steneleodes_gravidus_E30 Eleodes_esch_eschscholtzi_E23 Steneleodes_distinctus_E154 Eleodes_loretensis_TB15485 Eleodes_gracilis_E28 Eleodes_80_E37 Blaps_sp2_TB17180 Eleodes__Eleodes__loretensis_TB15983 Litheleodes_arcuatus_E140 Eleodes_eschscholtzi_E21 Eleodes_tenuipes_E17 Eleodes__Melaneleodes__debilis_TB15988 Steneleodes_angustus_E168 100 100 9 1 100 100 6 2 7 9 9 1 100 4 7 6 0 3 8 100 3 9 4 6 4 6 9 3 4 2 9 8 100 6 5 4 3 100 8 7 9 0 9 0 8 2 100 9 8 6 7 6 7 7 2 100 9 6 1 9 100 6 3 5 1 1 2 5 2 4 0 100 9 9 100 3 8 3 4 1 3 9 4 100 7 6 9 8 9 8 100 9 7 9 8 100 100 8 8 2 1 2 0 7 6 100 100 4 8 8 4 100 9 8 6 5 3 6 100 5 7 5 5 9 2 9 5 100 100 4 0 9 7 7 8 6 0 3 6 100 9 4 3 9 100 100 9 8 1 8 100 9 9 9 9 4 3 7 8 100 100 9 9 100 2 1 5 3 9 9 100 100 3 4 100 9 9 9 3 6 5 100 4 1 8 2 3 8 5 2 4 3 8 9 100 100 4 2 100 2 3 100 100 100 100 6 6 100 100 9 6 100 9 2 5 3 8 2 5 6 8 2 100 100 100 100 9 0 9 8 100 9 9 100 6 7 1 2 100 7 4 100 9 9 9 4 3 0 9 9 100 9 9 100 5 5 100 9 9 100 100 8 2 100 100 100 100 100 100 100 100
- 31. Why so many similar species? Generalist scavengers
- 32. Why so many similar species? Generalist scavengers Minimal genitalic differences
- 33. Why so many similar species? Extensive distributional overlap >200 taxa, 20,000 specimens (OSUC,MAJC)
- 34. Why so many similar species? Extensive distributional overlap Centers of diversity Pacific coast Intermountain region Northern Mexico Intermountain region? >200 taxa, 20,000 specimens (OSUC,MAJC)
- 35. Why so many similar species? Apparently mountain uplift and glaciation drove Amphidorini diversification Supported by Trogloderus Pleistocene diversification
- 36. When did Amphidorini originate? Eleodes spinipes macrura Champion, 1892
- 37. When did Amphidorini originate? Eleodes spinipes macrura Champion, 1892Blaps lethifera Marsham, 1802 © Stanislav Snäll Sister to Eurasian Blaptini Leach, 1815
- 38. When did Amphidorini originate? Smith et al. 2006 https://doi.org/10.1073/pnas.0511296103 Sister to Eurasian Blaptini Leach, 1815 • Blaptini ~55my old • Corresponds to mammal diversification across land bridges
- 39. Future directions Finish species-level revisions
- 40. Future directions Diversification analyses (but lack fossils) Finish species-level revisions
- 41. Future directions Historical biogeographic estimation for the tribe Diversification analyses (but lack fossils) Finish species-level revisions
- 42. Thank you! Funding: National Science Foundation (DEB–1258154, DEB–1754731) Visiting Scientist Fellowship CanaColl foundation Collaborators: Nico Franz Rolf Aalbu Rebecca Livingston Omron Blau Sydney Wuerker Travis Hitchner Lab members: Sangmi Lee Brian Reily Andrew Jansen Kathy Arguez Sal Anzaldo Ed Gilbert
Editor's Notes
- 430 mammals, 800 birds in the US
- 430 mammals, 800 birds in the US
- 430 mammals, 800 birds in the US