Johnston ESA 2014 Trogloderus Sand Dune Speciation
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Transcript of Johnston ESA 2014 Trogloderus Sand Dune Speciation
Sand Dune Speciation:Exploring the Evolutionary History of
Trogloderus LeConte
(Coleoptera: Tenebrionidae)
M. Andrew Johnston
Trogloderus LeConte 1879
• Erected as a monotypic genus
– 1 specimen from SW Idaho
• Flightless
• Psammophilic
– Eolian sand formations
• Western U.S. distribution
Trogloderus costatus LeConte
Dumont Dunes, CA
Olancha Dunes, CA
Trogloderus LeConte 1879
Tribe Amphidorini LeConte 1862
• North American distribution
• 6 genera, ~ 250 species
• Most in Eleodes Eschscholtz
Eleodes armatus LeConte
Taxonomic History
Revised by La Rivers, 1946
– 4 species subspecies
– 2 more described later
Trogloderus
costatus LeConte tuberculatus Blaisdell nevadus La Rivers vandykei La Rivers
costatus
ssp. ssp. ssp. ssp.
syn. mayhewi Pappsyn. pappi Kulzer
Taxonomic History
Revised by La Rivers, 1946
• Invoked orthogenetic evolution
• Hypothesized Trogloderus to be ancient
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?
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
Eleodes_hepburni
Eleodes_ruidus
Eleodes_scabriculus
Eleodes_innocens
Eleodes_hispilabris
Eleodes_nspTontNatBr
Eleodes_carb_omissus
Eleodes_subnitens
Eleodes_acuticaudus
Eleodes_impolitus
Eleodes_nigropilosus
Eleodes_spinipes_ventricosus
Blapstinus_fortis
Eleodes_giganteus
Trogloderus
Eleodes_veterator
Embaphion_muricatum
Eleodes_microps
Eleodes_mutilatus
Eleodes_sponsus
Eleodes_tenuipes
Centronopus_suppressus
Embaphion_glabrum
Neobaphion_planipennis
Eleodes_wynnei
Eleodes_rossi
Eleodes_striolatus
Embaphion_contusum
Eleodes_caudiferus
Eleodes_connatus
Eleodes_pilosus
Argoporis_cribrata
Eleodes_alutaceus
Eleodes_eschscholtzi
Eleodes_carb_carb
Eleodes_gravidus
Eleodes_tribulus
Eleodes_suturalis
Eleodes_gracilis
Eleodes_tricostatus
Eleodes_nigrinus
Rhinandrus_helopioides
Eleodes_clavicornis
Eleodes_debilis
Eleodes_osculans
Eleodes_spinolae
Eleodes_knullorum
Eleodes_insularis
Wattius_sp
Eleodes_snowii
Eleodes_wenzeli
Cymatothes_opacus
Eleodes_ponderosus
Lariversius
Eleodes_hogei
Eleodes_fusiformis
Eleodes_extricatus
Embaphion_depressum
Eleodes_obsc_sulcipennis
Eleodes_goryi
Nyctorinus_sp2
Eleodes_tenebrosus
Eleodes_carb_obsoletus
Eleodes_longicollis
Eleodes_dissimilis
Eleodes_esch_eschscholtzi
Eleodes_armatus
8 0
7 2
100
100
9 7
7 3
6 7
9 5
9 6
2 4
100
100
4 3
9 8
2 8
8 3
6 4
9 8
1 3
9 1
6 6
6 3
8 5
9 5
3 3
8 89 1
6 4
8
2 5
100
9 5
8 8
9 9
100
7 6
100
6 6
2 4
2 7
100
7 7
9 4
8 4
100
1 2
8 0
5 6
100
4 8
5 9
7 7
9 5
8 4
9 7
9 4
7 9
3 8
4 2
7 7
2 5
100
7 4
8 9
8 9
5 8
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
Eleodes_hepburni
Eleodes_ruidus
Eleodes_scabriculus
Eleodes_innocens
Eleodes_hispilabris
Eleodes_nspTontNatBr
Eleodes_carb_omissus
Eleodes_subnitens
Eleodes_acuticaudus
Eleodes_impolitus
Eleodes_nigropilosus
Eleodes_spinipes_ventricosus
Blapstinus_fortis
Eleodes_giganteus
Trogloderus
Eleodes_veterator
Embaphion_muricatum
Eleodes_microps
Eleodes_mutilatus
Eleodes_sponsus
Eleodes_tenuipes
Centronopus_suppressus
Embaphion_glabrum
Neobaphion_planipennis
Eleodes_wynnei
Eleodes_rossi
Eleodes_striolatus
Embaphion_contusum
Eleodes_caudiferus
Eleodes_connatus
Eleodes_pilosus
Argoporis_cribrata
Eleodes_alutaceus
Eleodes_eschscholtzi
Eleodes_carb_carb
Eleodes_gravidus
Eleodes_tribulus
Eleodes_suturalis
Eleodes_gracilis
Eleodes_tricostatus
Eleodes_nigrinus
Rhinandrus_helopioides
Eleodes_clavicornis
Eleodes_debilis
Eleodes_osculans
Eleodes_spinolae
Eleodes_knullorum
Eleodes_insularis
Wattius_sp
Eleodes_snowii
Eleodes_wenzeli
Cymatothes_opacus
Eleodes_ponderosus
Lariversius
Eleodes_hogei
Eleodes_fusiformis
Eleodes_extricatus
Embaphion_depressum
Eleodes_obsc_sulcipennis
Eleodes_goryi
Nyctorinus_sp2
Eleodes_tenebrosus
Eleodes_carb_obsoletus
Eleodes_longicollis
Eleodes_dissimilis
Eleodes_esch_eschscholtzi
Eleodes_armatus
8 0
7 2
100
100
9 7
7 3
6 7
9 5
9 6
2 4
100
100
4 3
9 8
2 8
8 3
6 4
9 8
1 3
9 1
6 6
6 3
8 5
9 5
3 3
8 89 1
6 4
8
2 5
100
9 5
8 8
9 9
100
7 6
100
6 6
2 4
2 7
100
7 7
9 4
8 4
100
1 2
8 0
5 6
100
4 8
5 9
7 7
9 5
8 4
9 7
9 4
7 9
3 8
4 2
7 7
2 5
100
7 4
8 9
8 9
5 8
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
Trogloderus Diversification
• Trogloderus recovered as
monophyletic
• Parent clade supported
• Sister relationship not well
supported
Eleodes_wynnei
Trogloderus_Hanksville
Lariversius_BigDune
Eleodes_esch_eschscholtzi
Eleodes_carb_obsoletus
Eleodes_dissimilis
Eleodes_caudiferus
Eleodes_hispilabris
Embaphion_glabrum
Argoporis_cribrata
Lariversius_tibialis
Eleodes_fusiformis
Eleodes_giganteus
Eleodes_extricatus
Eleodes_longicollis
Eleodes_impolitus
Eleodes_eschscholtzi
Eleodes_gravidus
Eleodes_barbatus
Eleodes_goryi
Eleodes_hogei
Eleodes_rossi
Embaphion_contusum
Eleodes_scabriculus
Eleodes_suturalis
Eleodes_alutaceus
Eleodes_innocens
Trogloderus_Chilchinbito
Eleodes_acuticaudus
Trogloderus_DumontDunes
Wattius_sp
Trogloderus_CrescentDunes
Trogloderus_BeaverDam
Lariversius_tibialis_WashoeCo
Eleodes_wenzeli
Eleodes_insularis
Blapstinus_fortis
Eleodes_nigrinus
Eleodes_tenebrosus
Eleodes_obsc_sulcipennis
Eleodes_mutilatus
Trogloderus_29palms
Eleodes_spinipes_ventricosus
Eleodes_veterator
Eleodes_spinolae
Trogloderus_VermCliffs
Eleodes_snowii
Trogloderus_MonoRetic
Centronopus_suppressus
Eleodes_microps
Eleodes_connatus
Eleodes_ponderosus
Eleodes_hepburni
Eleodes_clavicornis
Eleodes_pilosus
Eleodes_armatus
Nyctorinus_sp2
Eleodes_osculans
Eleodes_tenuipes
Embaphion_muricatum
Eleodes_carb_carb
Eleodes_tricostatus
Embaphion_depressum
Eleodes_ruidus
Cymatothes_opacus
Neobaphion_planipennis
Eleodes_nigropilosus
Eleodes_sponsus
Eleodes_gracilis
Trogloderus_PyramidLake
Trogloderus_Olancha
Trogloderus_ElMirage
Eleodes_tribulus
Eleodes_knullorum
Eleodes_debilis
Rhinandrus_helopioides
Eleodes_carb_omissus
Eleodes_striolatus
Eleodes_nspTontNatBr
Trogloderus_CoralPink
Trogloderus_MonoTuber
Eleodes_subnitens
8 8
100
3 8
8 3
9 4
9 6
2 5
100
1 3
7 4
9 5
5 8
100
100
100
100
100
2 8
9 3
4 2
100
7 7
100
9 1
3 4
9 8
8 4
9 5
7 2
6 4
9 1
100
5 6
8 8
2 4
100
7 3
2 1
6 7
8 0
100
2 7
100
4 3
6 3
8 9
3 3
7 6
7 4
100
100
8 5
100
8 8
8 9
6 6
8
9 4
8
4 8
6 4
9 5
2 5
9 9
2 4
6 6
100
8 0
7 7
9 7
7 9
8 4
1 2
7 7
100
4 2
9 8
9 7
5 9
9 5
Trogloderus Diversification
Close outgroups
• Mainly Eleodes (Blapylis)
• Dune Specialists
• High elevation species
– AZ, CA, Pacific NW
– Under-sampled
Eleodes_wynnei
Trogloderus_Hanksville
Lariversius_BigDune
Eleodes_esch_eschscholtzi
Eleodes_carb_obsoletus
Eleodes_dissimilis
Eleodes_caudiferus
Eleodes_hispilabris
Embaphion_glabrum
Argoporis_cribrata
Lariversius_tibialis
Eleodes_fusiformis
Eleodes_giganteus
Eleodes_extricatus
Eleodes_longicollis
Eleodes_impolitus
Eleodes_eschscholtzi
Eleodes_gravidus
Eleodes_barbatus
Eleodes_goryi
Eleodes_hogei
Eleodes_rossi
Embaphion_contusum
Eleodes_scabriculus
Eleodes_suturalis
Eleodes_alutaceus
Eleodes_innocens
Trogloderus_Chilchinbito
Eleodes_acuticaudus
Trogloderus_DumontDunes
Wattius_sp
Trogloderus_CrescentDunes
Trogloderus_BeaverDam
Lariversius_tibialis_WashoeCo
Eleodes_wenzeli
Eleodes_insularis
Blapstinus_fortis
Eleodes_nigrinus
Eleodes_tenebrosus
Eleodes_obsc_sulcipennis
Eleodes_mutilatus
Trogloderus_29palms
Eleodes_spinipes_ventricosus
Eleodes_veterator
Eleodes_spinolae
Trogloderus_VermCliffs
Eleodes_snowii
Trogloderus_MonoRetic
Centronopus_suppressus
Eleodes_microps
Eleodes_connatus
Eleodes_ponderosus
Eleodes_hepburni
Eleodes_clavicornis
Eleodes_pilosus
Eleodes_armatus
Nyctorinus_sp2
Eleodes_osculans
Eleodes_tenuipes
Embaphion_muricatum
Eleodes_carb_carb
Eleodes_tricostatus
Embaphion_depressum
Eleodes_ruidus
Cymatothes_opacus
Neobaphion_planipennis
Eleodes_nigropilosus
Eleodes_sponsus
Eleodes_gracilis
Trogloderus_PyramidLake
Trogloderus_Olancha
Trogloderus_ElMirage
Eleodes_tribulus
Eleodes_knullorum
Eleodes_debilis
Rhinandrus_helopioides
Eleodes_carb_omissus
Eleodes_striolatus
Eleodes_nspTontNatBr
Trogloderus_CoralPink
Trogloderus_MonoTuber
Eleodes_subnitens
8 8
100
3 8
8 3
9 4
9 6
2 5
100
1 3
7 4
9 5
5 8
100
100
100
100
100
2 8
9 3
4 2
100
7 7
100
9 1
3 4
9 8
8 4
9 5
7 2
6 4
9 1
100
5 6
8 8
2 4
100
7 3
2 1
6 7
8 0
100
2 7
100
4 3
6 3
8 9
3 3
7 6
7 4
100
100
8 5
100
8 8
8 9
6 6
8
9 4
8
4 8
6 4
9 5
2 5
9 9
2 4
6 6
100
8 0
7 7
9 7
7 9
8 4
1 2
7 7
100
4 2
9 8
9 7
5 9
9 5
Photo by Kojun Kanda
Eleodes barbatus Whickham Lariversius tibialis BlaisdellEleodes snowii Blaisdell
Trogloderus Diversification
• Molecular phylogeny supports
and explains the 7 identified
morphotypes
• Pronotal Sculpturing
Eleodes_wynnei
Trogloderus_Hanksville
Lariversius_BigDune
Eleodes_esch_eschscholtzi
Eleodes_carb_obsoletus
Eleodes_dissimilis
Eleodes_caudiferus
Eleodes_hispilabris
Embaphion_glabrum
Argoporis_cribrata
Lariversius_tibialis
Eleodes_fusiformis
Eleodes_giganteus
Eleodes_extricatus
Eleodes_longicollis
Eleodes_impolitus
Eleodes_eschscholtzi
Eleodes_gravidus
Eleodes_barbatus
Eleodes_goryi
Eleodes_hogei
Eleodes_rossi
Embaphion_contusum
Eleodes_scabriculus
Eleodes_suturalis
Eleodes_alutaceus
Eleodes_innocens
Trogloderus_Chilchinbito
Eleodes_acuticaudus
Trogloderus_DumontDunes
Wattius_sp
Trogloderus_CrescentDunes
Trogloderus_BeaverDam
Lariversius_tibialis_WashoeCo
Eleodes_wenzeli
Eleodes_insularis
Blapstinus_fortis
Eleodes_nigrinus
Eleodes_tenebrosus
Eleodes_obsc_sulcipennis
Eleodes_mutilatus
Trogloderus_29palms
Eleodes_spinipes_ventricosus
Eleodes_veterator
Eleodes_spinolae
Trogloderus_VermCliffs
Eleodes_snowii
Trogloderus_MonoRetic
Centronopus_suppressus
Eleodes_microps
Eleodes_connatus
Eleodes_ponderosus
Eleodes_hepburni
Eleodes_clavicornis
Eleodes_pilosus
Eleodes_armatus
Nyctorinus_sp2
Eleodes_osculans
Eleodes_tenuipes
Embaphion_muricatum
Eleodes_carb_carb
Eleodes_tricostatus
Embaphion_depressum
Eleodes_ruidus
Cymatothes_opacus
Neobaphion_planipennis
Eleodes_nigropilosus
Eleodes_sponsus
Eleodes_gracilis
Trogloderus_PyramidLake
Trogloderus_Olancha
Trogloderus_ElMirage
Eleodes_tribulus
Eleodes_knullorum
Eleodes_debilis
Rhinandrus_helopioides
Eleodes_carb_omissus
Eleodes_striolatus
Eleodes_nspTontNatBr
Trogloderus_CoralPink
Trogloderus_MonoTuber
Eleodes_subnitens
8 8
100
3 8
8 3
9 4
9 6
2 5
100
1 3
7 4
9 5
5 8
100
100
100
100
100
2 8
9 3
4 2
100
7 7
100
9 1
3 4
9 8
8 4
9 5
7 2
6 4
9 1
100
5 6
8 8
2 4
100
7 3
2 1
6 7
8 0
100
2 7
100
4 3
6 3
8 9
3 3
7 6
7 4
100
100
8 5
100
8 8
8 9
6 6
8
9 4
8
4 8
6 4
9 5
2 5
9 9
2 4
6 6
100
8 0
7 7
9 7
7 9
8 4
1 2
7 7
100
4 2
9 8
9 7
5 9
9 5
tuber
cula
tere
ticu
late
Sympatric
5% sequence
divergence
Trogloderus Distribution
Eleodes_wynnei
Trogloderus_Hanksville
Lariversius_BigDune
Eleodes_esch_eschscholtzi
Eleodes_carb_obsoletus
Eleodes_dissimilis
Eleodes_caudiferus
Eleodes_hispilabris
Embaphion_glabrum
Argoporis_cribrata
Lariversius_tibialis
Eleodes_fusiformis
Eleodes_giganteus
Eleodes_extricatus
Eleodes_longicollis
Eleodes_impolitus
Eleodes_eschscholtzi
Eleodes_gravidus
Eleodes_barbatus
Eleodes_goryi
Eleodes_hogei
Eleodes_rossi
Embaphion_contusum
Eleodes_scabriculus
Eleodes_suturalis
Eleodes_alutaceus
Eleodes_innocens
Trogloderus_Chilchinbito
Eleodes_acuticaudus
Trogloderus_DumontDunes
Wattius_sp
Trogloderus_CrescentDunes
Trogloderus_BeaverDam
Lariversius_tibialis_WashoeCo
Eleodes_wenzeli
Eleodes_insularis
Blapstinus_fortis
Eleodes_nigrinus
Eleodes_tenebrosus
Eleodes_obsc_sulcipennis
Eleodes_mutilatus
Trogloderus_29palms
Eleodes_spinipes_ventricosus
Eleodes_veterator
Eleodes_spinolae
Trogloderus_VermCliffs
Eleodes_snowii
Trogloderus_MonoRetic
Centronopus_suppressus
Eleodes_microps
Eleodes_connatus
Eleodes_ponderosus
Eleodes_hepburni
Eleodes_clavicornis
Eleodes_pilosus
Eleodes_armatus
Nyctorinus_sp2
Eleodes_osculans
Eleodes_tenuipes
Embaphion_muricatum
Eleodes_carb_carb
Eleodes_tricostatus
Embaphion_depressum
Eleodes_ruidus
Cymatothes_opacus
Neobaphion_planipennis
Eleodes_nigropilosus
Eleodes_sponsus
Eleodes_gracilis
Trogloderus_PyramidLake
Trogloderus_Olancha
Trogloderus_ElMirage
Eleodes_tribulus
Eleodes_knullorum
Eleodes_debilis
Rhinandrus_helopioides
Eleodes_carb_omissus
Eleodes_striolatus
Eleodes_nspTontNatBr
Trogloderus_CoralPink
Trogloderus_MonoTuber
Eleodes_subnitens
8 8
100
3 8
8 3
9 4
9 6
2 5
100
1 3
7 4
9 5
5 8
100
100
100
100
100
2 8
9 3
4 2
100
7 7
100
9 1
3 4
9 8
8 4
9 5
7 2
6 4
9 1
100
5 6
8 8
2 4
100
7 3
2 1
6 7
8 0
100
2 7
100
4 3
6 3
8 9
3 3
7 6
7 4
100
100
8 5
100
8 8
8 9
6 6
8
9 4
8
4 8
6 4
9 5
2 5
9 9
2 4
6 6
100
8 0
7 7
9 7
7 9
8 4
1 2
7 7
100
4 2
9 8
9 7
5 9
9 5
0.1% within-group,
2% between-group
sequence divergence
T. “warneri” sp nov
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
Trogloderus Distribution
T. “warneri” sp nov• Gene flow inhibited by
geographic barriers
– Wasatch Mountains
– Colorado River
Trogloderus Distribution
Eleodes_wynnei
Trogloderus_Hanksville
Lariversius_BigDune
Eleodes_esch_eschscholtzi
Eleodes_carb_obsoletus
Eleodes_dissimilis
Eleodes_caudiferus
Eleodes_hispilabris
Embaphion_glabrum
Argoporis_cribrata
Lariversius_tibialis
Eleodes_fusiformis
Eleodes_giganteus
Eleodes_extricatus
Eleodes_longicollis
Eleodes_impolitus
Eleodes_eschscholtzi
Eleodes_gravidus
Eleodes_barbatus
Eleodes_goryi
Eleodes_hogei
Eleodes_rossi
Embaphion_contusum
Eleodes_scabriculus
Eleodes_suturalis
Eleodes_alutaceus
Eleodes_innocens
Trogloderus_Chilchinbito
Eleodes_acuticaudus
Trogloderus_DumontDunes
Wattius_sp
Trogloderus_CrescentDunes
Trogloderus_BeaverDam
Lariversius_tibialis_WashoeCo
Eleodes_wenzeli
Eleodes_insularis
Blapstinus_fortis
Eleodes_nigrinus
Eleodes_tenebrosus
Eleodes_obsc_sulcipennis
Eleodes_mutilatus
Trogloderus_29palms
Eleodes_spinipes_ventricosus
Eleodes_veterator
Eleodes_spinolae
Trogloderus_VermCliffs
Eleodes_snowii
Trogloderus_MonoRetic
Centronopus_suppressus
Eleodes_microps
Eleodes_connatus
Eleodes_ponderosus
Eleodes_hepburni
Eleodes_clavicornis
Eleodes_pilosus
Eleodes_armatus
Nyctorinus_sp2
Eleodes_osculans
Eleodes_tenuipes
Embaphion_muricatum
Eleodes_carb_carb
Eleodes_tricostatus
Embaphion_depressum
Eleodes_ruidus
Cymatothes_opacus
Neobaphion_planipennis
Eleodes_nigropilosus
Eleodes_sponsus
Eleodes_gracilis
Trogloderus_PyramidLake
Trogloderus_Olancha
Trogloderus_ElMirage
Eleodes_tribulus
Eleodes_knullorum
Eleodes_debilis
Rhinandrus_helopioides
Eleodes_carb_omissus
Eleodes_striolatus
Eleodes_nspTontNatBr
Trogloderus_CoralPink
Trogloderus_MonoTuber
Eleodes_subnitens
8 8
100
3 8
8 3
9 4
9 6
2 5
100
1 3
7 4
9 5
5 8
100
100
100
100
100
2 8
9 3
4 2
100
7 7
100
9 1
3 4
9 8
8 4
9 5
7 2
6 4
9 1
100
5 6
8 8
2 4
100
7 3
2 1
6 7
8 0
100
2 7
100
4 3
6 3
8 9
3 3
7 6
7 4
100
100
8 5
100
8 8
8 9
6 6
8
9 4
8
4 8
6 4
9 5
2 5
9 9
2 4
6 6
100
8 0
7 7
9 7
7 9
8 4
1 2
7 7
100
4 2
9 8
9 7
5 9
9 5
Overlapping ranges,
4% sequence divergence
Different male genitalia
T. nevadus nevadus La
Rivers
T. vandykei La Rivers
T. nevadus “aalbui” sp nov
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
Trogloderus Distribution
Unplaced Species
• T. costatus LeConte
• T. tuberculatus Blaisdell
Results
• Does current taxonomy capture known
diversity?
– 5 species, 1 currently undescribed
– 2 undescribed subspecies
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
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
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
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 CreditsKojun Kanda, Bugguide.net
Aaron Smith, tenebrioniDBase.org
ETOPO1, NOAA