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