Sand Dune Speciation: Exploring the Evolutionary History of Trogloderus LeConte
- 1. Sand Dune Speciation: Exploring the Evolutionary History of Trogloderus LeConte (Coleoptera: Tenebrionidae) M. Andrew Johnston
- 2. Trogloderus LeConte 1879 • Erected as a monotypic genus – 1 specimen from SW Idaho • Flightless • Psammophilic Trogloderus costatus LeConte – Eolian sand formations • Western U.S. distribution Dumont Dunes, CA Olancha Dunes, CA
- 3. Trogloderus LeConte 1879 Tribe Amphidorini LeConte 1862 • North American distribution • 6 genera, ~ 250 species • Most in Eleodes Eschscholtz Eleodes armatus LeConte
- 4. Taxonomic History Revised by La Rivers, 1946 Trogloderus costatus costatus LeConte tuberculatus Blaisdell nevadus La Rivers vandykei La Rivers ssp. ssp. ssp. ssp. syn. pappi Kulzer syn. mayhewi Papp – 4 species subspecies – 2 more described later
- 5. Taxonomic History Revised by La Rivers, 1946 • Invoked orthogenetic evolution • Hypothesized Trogloderus to be ancient
- 6. Research Questions • Does the current taxonomy capture actual diversity? • When and where did Trogloderus originate? • What are the patterns and processes of diversification across sand dunes?
- 7. Research Methods • Morphological analysis – 1,860 Trogloderus specimens – Evaluated morphotypes and their distributions • Molecular analysis – Phylogenetic reconstruction of Amphidorini • 77 taxa, 7 gene fragments – 3 nuclear, 4 mitochondrial • Aligned in MUSCLE • Partitioned by PartitionFinder • Analyzed by RAxML, MrBayes and BEAST
- 8. Wattius_sp Argoporis_cribrata Rhinandrus_helopioides Nyctorinus_sp2 Cymatothes_opacus Centronopus_suppressus Blapstinus_fortis Neobaphion_planipennis Embaphion_muricatum Embaphion_contusum Embaphion_depressum Embaphion_glabrum Eleodes_osculans Eleodes_nigropilosus Eleodes_wynnei Eleodes_microps Eleodes_nigrinus Eleodes_nspTontNatBr Eleodes_dissimilis Eleodes_ponderosus Eleodes_innocens Eleodes_mutilatus Eleodes_impolitus Eleodes_hepburni Eleodes_gravidus Eleodes_ruidus Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Lariversius Trogloderus Eleodes_spinipes_ventricosus Eleodes_debilis Eleodes_hogei Eleodes_connatus Eleodes_tribulus Eleodes_caudiferus Eleodes_pilosus Eleodes_veterator Eleodes_alutaceus Eleodes_wenzeli Eleodes_tricostatus Eleodes_carb_carb Eleodes_carb_omissus Eleodes_carb_obsoletus Eleodes_acuticaudus Eleodes_obsc_sulcipennis Eleodes_suturalis Eleodes_esch_eschscholtzi Eleodes_hispilabris Eleodes_sponsus Eleodes_gracilis Eleodes_tenuipes Eleodes_armatus Eleodes_rossi Eleodes_eschscholtzi Eleodes_subnitens Eleodes_spinolae Eleodes_knullorum Eleodes_insularis Eleodes_goryi Eleodes_fusiformis Eleodes_striolatus Eleodes_extricatus 80 72 100 100 99 97 73 95 84 67 25 95 96 24 100 100 43 98 28 83 64 98 13 91 66 63 85 95 33 88 91 64 8 25 100 95 88 100 76 100 66 24 27 100 77 94 84 100 12 56 80 100 48 59 77 97 94 79 38 42 77 100 74 89 89 58 8 Amphidorini Phylogeny • All analyses highly congruent • Eleodes paraphyletic – Trogloderus LeConte – Lariversius Blaisdell ML Reconstruction Showing Bootstrap Support Photo by Kojun Kanda Lariversius tibialis Blaisdell
- 9. Wattius_sp Argoporis_cribrata Rhinandrus_helopioides Nyctorinus_sp2 Cymatothes_opacus Centronopus_suppressus Blapstinus_fortis Neobaphion_planipennis Embaphion_muricatum Embaphion_contusum Embaphion_depressum Embaphion_glabrum Eleodes_osculans Eleodes_nigropilosus Eleodes_wynnei Eleodes_microps Eleodes_nigrinus Eleodes_nspTontNatBr Eleodes_dissimilis Eleodes_ponderosus Eleodes_innocens Eleodes_mutilatus Eleodes_impolitus Eleodes_hepburni Eleodes_gravidus Eleodes_ruidus Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Lariversius Trogloderus Eleodes_spinipes_ventricosus Eleodes_debilis Eleodes_hogei Eleodes_connatus Eleodes_tribulus Eleodes_caudiferus Eleodes_pilosus Eleodes_veterator Eleodes_alutaceus Eleodes_wenzeli Eleodes_tricostatus Eleodes_carb_carb Eleodes_carb_omissus Eleodes_carb_obsoletus Eleodes_acuticaudus Eleodes_obsc_sulcipennis Eleodes_suturalis Eleodes_esch_eschscholtzi Eleodes_hispilabris Eleodes_sponsus Eleodes_gracilis Eleodes_tenuipes Eleodes_armatus Eleodes_rossi Eleodes_eschscholtzi Eleodes_subnitens Eleodes_spinolae Eleodes_knullorum Eleodes_insularis Eleodes_goryi Eleodes_fusiformis Eleodes_striolatus Eleodes_extricatus 80 72 100 100 99 97 73 95 84 67 25 95 96 24 100 100 43 98 28 83 64 98 13 91 66 63 85 95 33 88 91 64 8 25 100 95 88 100 76 100 66 24 27 100 77 94 84 100 12 56 80 100 48 59 77 97 94 79 38 42 77 100 74 89 89 58 8 Amphidorini Phylogeny • Dune specialization is common – Found in at least 7 lineages – Ancestral state unclear ML Reconstruction Showing Bootstrap Support Photo by Kojun Kanda Lariversius tibialis Blaisdell
- 10. Trogloderus Diversification • Trogloderus recovered as monophyletic • Parent clade supported • Sister relationship not well supported Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Eleodes_barbatus Lariversius_tibialis_WashoeCo Lariversius_tibialis Lariversius_BigDune Trogloderus_Chilchinbito 100 Trogloderus_Hanksville Trogloderus_VermCliffs Trogloderus_CoralPink Trogloderus_MonoTuber 100 Trogloderus_BeaverDam Trogloderus_29palms Trogloderus_Olancha Trogloderus_PyramidLake Trogloderus_ElMirage Trogloderus_DumontDunes Trogloderus_CrescentDunes Trogloderus_MonoRetic Eleodes_spinipes_ventricosus 25 74 100 13 74 77 100 93 42 100 34 91 88 21 80 27 100 100 94 84
- 11. Trogloderus Diversification Close outgroups • Mainly Eleodes (Blapylis) • Dune Specialists • High elevation species – AZ, CA, Pacific NW – Under-sampled Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Eleodes_barbatus Lariversius_tibialis_WashoeCo Lariversius_tibialis Lariversius_BigDune Trogloderus_Chilchinbito 100 Trogloderus_Hanksville Trogloderus_VermCliffs Trogloderus_CoralPink Trogloderus_MonoTuber 100 Trogloderus_BeaverDam Trogloderus_29palms Trogloderus_Olancha Trogloderus_PyramidLake Trogloderus_ElMirage Trogloderus_DumontDunes Trogloderus_CrescentDunes Trogloderus_MonoRetic Eleodes_spinipes_ventricosus 25 74 100 13 74 77 100 93 42 100 34 91 88 21 80 27 100 100 94 84 Photo by Kojun Kanda Eleodes barbatusWhickham Eleodes snowii BlLaaisrdiveellrsius tibialis Blaisdell
- 12. Trogloderus Diversification • Molecular phylogeny supports and explains the 7 identified morphotypes • Pronotal Sculpturing Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Eleodes_barbatus Lariversius_tibialis_WashoeCo Lariversius_tibialis Lariversius_BigDune Trogloderus_Chilchinbito 100 Trogloderus_Hanksville Trogloderus_VermCliffs Trogloderus_CoralPink Trogloderus_MonoTuber 100 Trogloderus_BeaverDam Trogloderus_29palms Trogloderus_Olancha Trogloderus_PyramidLake Trogloderus_ElMirage Trogloderus_DumontDunes Trogloderus_CrescentDunes Trogloderus_MonoRetic Eleodes_spinipes_ventricosus 25 74 100 13 74 77 100 93 42 100 34 91 88 21 80 27 100 100 94 84 tuberculate reticulate Sympatric 5% sequence divergence
- 13. Trogloderus Distribution Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Eleodes_barbatus Lariversius_tibialis_WashoeCo Lariversius_tibialis Lariversius_BigDune Trogloderus_Chilchinbito 100 Trogloderus_Hanksville Trogloderus_VermCliffs Trogloderus_CoralPink Trogloderus_MonoTuber 100 Trogloderus_BeaverDam Trogloderus_29palms Trogloderus_Olancha Trogloderus_PyramidLake Trogloderus_ElMirage Trogloderus_DumontDunes Trogloderus_CrescentDunes Trogloderus_MonoRetic Eleodes_spinipes_ventricosus 25 74 100 13 74 77 100 93 42 100 34 91 88 21 80 27 100 100 94 84 0.1% within-group, 2% between-group sequence divergence T. “warneri” sp nov
- 14. Trogloderus Distribution T. “warneri” sp nov Great Basin Colorado Plateau Costae tuberculate Rugose intervals Costae and intervals smooth Transitional population found along Northern bank of Lake Powell
- 15. Trogloderus Distribution T. “warneri” sp nov • Gene flow inhibited by geographic barriers – Wasatch Mountains – Colorado River
- 16. Trogloderus Distribution Eleodes_giganteus Eleodes_longicollis Eleodes_scabriculus Eleodes_snowii Eleodes_clavicornis Eleodes_tenebrosus Eleodes_barbatus Lariversius_tibialis_WashoeCo Lariversius_tibialis Lariversius_BigDune Trogloderus_Chilchinbito 100 Trogloderus_Hanksville Trogloderus_VermCliffs Trogloderus_CoralPink Trogloderus_MonoTuber 100 Trogloderus_BeaverDam Trogloderus_29palms Trogloderus_Olancha Trogloderus_PyramidLake Trogloderus_ElMirage Trogloderus_DumontDunes Trogloderus_CrescentDunes Trogloderus_MonoRetic Eleodes_spinipes_ventricosus 25 74 100 13 74 77 100 93 42 100 34 91 88 21 80 27 100 100 94 84 Overlapping ranges, 4% sequence divergence Different male genitalia T. nevadus nevadus La Rivers T. vandykei La Rivers T. nevadus “aalbui” sp nov
- 17. Trogloderus Distribution T. nevadus nevadus La Rivers T. vandykei La Rivers T. nevadus “aalbui” sp nov Great Basin dune formation • Late pleistocene 1-3 ma • Sand derived from dried pluvial lakes Owens Valley • Sand from Owens river • 10-35 ka Mojave Desert sand dunes • Sand from Mojave river • 10-35 ka
- 18. Trogloderus Distribution Unplaced Species • T. costatus LeConte • T. tuberculatus Blaisdell
- 19. Results • Does current taxonomy capture known diversity? – 5 species, 1 currently undescribed – 2 undescribed subspecies
- 20. Results • Does current taxonomy capture known diversity? • When and where did Trogloderus originate? – Likely a young lineage, perhaps with roots in Pleistocene – Area of origin unclear, perhaps in the Great Basin
- 21. Results • Does current taxonomy capture known diversity? • When and where did Trogloderus originate? • What are the patterns and processes of diversification? – Following recent sand formations, 10,000-35,000 years ago – Mountains and rivers that stop sand, stop gene flow
- 22. Future Directions • Finalize taxonomic revision of the genus • Increase taxon and geographic sampling for phylogenetic analyses • Develop calibrated dating analyses • Compare Trogloderus to other dune specialist groups
- 23. Acknowledgements Funding NSF ARTS award DEB-1258154 Specimens and Expertise Bill Warner Phoenix, AZ Rolf Aalbu CASC Warren Steiner USNM Lee Herman AMNH Kojun Kanda OSU Research Group Nico Franz Aaron Smith Sangmi Lee Sal Anzaldo Guanyang Zhang Chris Henny Andrew Jansen Omron Blauo Rebecca Livingston Photo Credits Kojun Kanda, Bugguide.net Aaron Smith, tenebrioniDBase.org ETOPO1, NOAA
Editor's Notes
- #2: ----- Meeting Notes (11/6/14 12:06) ----- TENEBRIONIDAE!!!!
- #5: ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
- #6: ----- Meeting Notes (11/6/14 12:06) ----- Explain orthogenic, or do not use
- #7: ----- Meeting Notes (11/13/14 12:01) ----- Add Sand Dunes to this slide.
- #9: ----- Meeting Notes (11/6/14 12:06) ----- Insert research objectives
- #10: ----- Meeting Notes (11/6/14 12:06) ----- Insert research objectives