A fistful of Astragalus: : incipient speciation in the American West? Brian J. Knaus Oregon State University Botany & Plant Pathology

Family Scientific Name Infra-rank rank count Fabaceae Astragalus lentiginosus var. 35 Polygonaceae Eriogonum umbellatum var. 30 Asteraceae Ericameria nauseosa ssp. & var. Asteraceae Hymenopappus filifolius var. 3 Malvaceae Sidalcea malviflora ssp. 3 Polygonaceae Eriogonum nudum var. 3 Asteraceae Ericameria parryi var. Asteraceae Eriophyllum lanatum var. Brassicaceae Lepidium montanum var. Asteraceae Achillea millefolium var. Caryophyllaceae Arenaria congesta var. Fabaceae Trifolium longipes ssp. Rosaceae Potentilla glandulosa ssp. USDA, NRCS. 006. The PLANTS database (http://plants.usda.gov, Nov. 006), National Plant Data Center, Baton Rogue, LA 70874-4490 USA.

A history of A. lentiginosus nomenclature Author Jones, M.E. Rydberg, P.A. Barneby, R.C. Barneby, R.C. Isely, D. NRCS, PLANTS Year 93 99 945 964 998 006 Taxa 8 36 40 36 40 35 Welsh, S.L. 007 4 Includes Tium sect. Palantia and Cystium sects., and Diphysa.

A. lentiginosus Douglas ex Hooker 83 Cystium Steven Coulteri A.l. var. ambiguus Barneby 964 A.l. var. australis Barneby 945 A.l. var. borreganus M.E. Jones 898 A.l. var. coachellae Barneby ap. Shreve & Wiggins 964 A.l. var. fremontii (A. Gray 857) S. Watson 87 A.l. var. kennedyi (Rydb. 99) Barneby 945 A.l. var. micans Barneby 956 A.l. var. nigricalycis M.E. Jones 895 A.l. var. stramineus (Rydb. 99) Barneby 945 A.l. var. variabilis Barneby 945 A.l. var. vitreus Barneby 945 A.l. var. yuccanus M.E. Jones 898 Cystium A.l. var. albifolius M.E. Jones 93 A.l. var. antonius Barneby 945 A.l. var. floribundus A. Gray 865 A.l. var. ineptus (A. Gray 864) M.E. Jones 93 A.l. var. kernensis (Jepson 95) Barneby 945 A.l. var. lentiginosus A.l. var. salinus (Howell 893) Barneby 945 A.l. var. scorpionis M.E. Jones 93 A.l. var. semotus Jepson 936 A.l. var. sierrae M.E. Jones 93 Cystium Diphysa A.l. var. araneosus (Sheldon 894) Barneby 945 A.l. var. chartaceus M.E. Jones 895 A.l. var. diphysus (A. Gray 849) M.E. Jones 895 A.l. var. higginsii S.L. Welsh & K.H. Thorne 98 A.l. var. idriensis M.E. Jones 90 A.l. var. latus (M.E. Jones 893) M.E. Jones 93 A.l. var. oropedii Barneby 945 A.l. var. piscinensis Barneby 977 A.l. var. pohlii S.L. Welsh & Barneby 98 A.l. var. sesquimetralis (Rydb. 99) Barneby 945 Tium Medic. Palantia A.l. var. maricopae Barneby 945 A.l. var. mokiacesis (A. Gray 878) M.E. Jones 93 A.l. var. palans (M.E. Jones 893) M.E. Jones 898 A.l. var. trumbellensis Welsh & Atwood 00 A.l. var. ursinus (A. Gray 878) Barneby 945 A.l. var. wilsonii (Greene 897) Barneby 945 38 varieties!

Distribution of A. lentiginosus

Variety Rydberg s Sections Distribution Barneby's Sample Barneby's Specimens Current Sample A. l. var. borreganus 3 4 9 A. l. var. coachellae 34 0 A. l. var. kennedyi 3 3 9 A. l. var. variabilis 4 9 A. l. var. araneosus Diphysa 39 8 0 A. l. var. chartaceus Diphysa 55 8 0 A. l. var. diphysus Diphysa 66 0 6 A. l. var. floribundus 6 4 A. l. var. fremontii 0 4 A. l. var. ineptus 3 A. l. var. lentiginosus 73 3 3 A. l. var. salinus 80 0 0 A. l. var. scorpionis 6 0 A. l. var. palans Palantia 40 9 0 4 varieties 739 85 44

Linear Model: Principle components as a function of climate Estimate Std. Error t value Pr(> t ) (Intercept) 8.7839 0.843 0.66 <e-6 data_lm$iv -.5999 0.439 -. <e-6 Residual standard error:.78 on 96 degrees of freedom Multiple R-Squared: 0.3869, Adjusted R-squared: 0.3838 F-statistic: 3.7 on and 96 DF, p-value: <.e-6 Estimate Std. Error t value Pr(> t ) (Intercept) -.34373 0.833-5.883.0e-08 data_lm$iv 0.00809 0.00573 6.87.08e-0 Residual standard error:.375 on 74 degrees of freedom Multiple R-Squared: 0.35, Adjusted R-squared: 0.09 F-statistic: 47.3 on and 74 DF, p-value:.08e-0

Discriminant Function Analysis to Distinguish Barneby s varieties. Color represents Rydberg s sections. DF Only Morphology Morphology with Latitude and Longitude DF & 3

A.l. var. salinus A.l. var. australis A.l. var. lentiginosus A.l. var. coachellae A.l. var. borreganus A.l. var. variabilis Green Grey A.l. var. lentiginosus A.l. var. australis A.l. var. borreganus White Purple

DNA Microsatellites a.k.a. simple sequence repeats (SSRs). Polymerases aren t good at counting, leading to length polymorphism. 300 bp 00 bp trntl839f ccmp0 ccmp5

Variety A. l. var. australis A. l. var. borreganus A. l. var. coachellae A. l. var. fremontii A. l. var. kennedyi A. l. var. micans A. l. var. nigricalycis A. l. var. stramineus A. l. var. variabilis A. l. var. vitreus A. l. var. yuccanus A. l. var. araneosus A. l. var. chartaceus A. l. var. idriensis A. l. var. piscinensis A. l. var. sesquimetralis A. l. var. albifolius A. l. var. antonius A. l. var. floribundus A. l. var. ineptus A. l. var. kernensis A. l. var. lentiginosus A. l. var. salinus A. l. var. scorpionis A. l. var. semotus A. l. var. sierrae A. l. var. maricopae A. l. var. mokiacensis A. l. var. palans A. l. var. trumbellensis A. l. var. ursinus A. l. var. wilsonii Rydbergian Section Diphysa Diphysa Diphysa Diphysa Diphysa Palantia Palantia Palantia Palantia Palantia Palantia Distribution Barneby's Sample 3 3 34 0 3 5 56 9 4 6 6 39 55 4 NA 6 8 6 3 8 73 80 6 4 8 5 5 40 NA 6 Barneby's Specimens 5 4 4 3 0 9 3 8 8 NA 0 3 0 0 3 0 0 0 0 9 NA 0 CpSSR pops 3 8 8 3 4 9 3 varieties 4 sections 3 varieties 68 populations 73 individuals 906 94 68

Geneland k = 7 (Only A. lentiginosus)

Incongruence of molecular markers and taxonomy?! The taxa are bad. Assumptions are bad. The marker is bad (e.g., mutation rate too high). Soil seed bank largely overlapping generations. Incomplete lineage sorting due to: Large ancestral effective population size. Recent divergence.

A fistful of Astragalus: : incipient speciation in the American West? Brian J. Knaus Oregon State University Botany & Plant Pathology

Dr. Rich Cronn (USDA FS) Dr. Matt Horning Rebecca Huot Sue Huber Angie Rodriguez Nancy Mandel Dr. Randy Johnson Chris Poklemba Dr. Brad St.Clair Dr. Nahla Bassil (USDA ARS) Wambui Njuguna April Nyberg Dr. Aaron Liston (OSU BPP) Dr. Richard Halse Drs. Ken & Henny Chambers Dr. John Syring Dr. Ann Wilyard Paul Severns Jason Alexander (leaf tissue) Dr. Bruce McCune (OSU BPP) Dr. Sarah Jovan Heather Lintz Dr. Emily Holt Heather Root Funding: NPSO field research grant Nevada Native Plant Society USDA FS PNW Research Station Hardman Foundation Bonnie C. Templeton Award