Publications - Papers available here
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Hinrichs, D.M., C.M. Patterson, A. Turcu, S.D. Holt Jr., S.G. Innes, N.J. Kooyers. In Review. Rapid adaptation follows experimental assisted gene flow in subset of annual monkeyflower populations. bioRxiv https://doi.org/10.64898/2025.12.02.689336
Gaspard, L.F., E. Harb, S.J. Holt Jr., C.M. Patterson, N.J. Kooyers. Accepted. Interactions between maternal provisioning and natural selection on seed mass fluctuate across heat waves. American Journal of Botany
Albano, L.J., C.M. Patterson, E.J. Kidder, N.J. Kooyers, M.T.J. Johnson. 2026. A field experiment assessing the roles of drought, herbivory, and local climate on cyanogenesis cline formation and local adaptation in Trifolium repens. Journal of Evolutionary Biology 39(4): 535-551
Albano, L.J., C.C. Bastias, A. Estarague, B.T. Hendrickson, S.G. Innes, N. King, C.M. Patterson, A. Tudoran, F. Vasseur, A. Puentes, C. Violle, N.J. Kooyers, M.T.J. Johnson. 2026. A transcontinental experiment elucidates (mal)adaptation of a cosmopolitan plant to climate in space and time. Ecological Monographs 96(1): e70043.
Innes, S.I., L.M. Holeski, N.J. Kooyers. 2025. Ontogenetic and geographic phytochemical variation in Mimulus moschatus, a perennial monkeyflower. Journal of Chemical Ecology. 51 (106): 1-12.
Kooyers, N. J., M.A. Genung, S.G. Innes, A.Turcu, D.M. Hinrichs, B.J. LeBlanc, and C.M. Patterson. 2025. Heat waves decrease fitness and alter maternal provisioning in natural populations of Mimulus guttatus. American Journal of Botany 112(8): e70087.
Battlay P., B.T. Hendrickson, J.I. Mendez-Reneau, J.S. Santangelo, L.J. Albano, J. Wilson, A.E. Caizergues, N. King, A. Puentes, A. Tudoran, C. Violle, F. Vasseur, C.M. Patterson, M. Foster, C. Stamps, S.G. Innes, R. Allio, F. Angeoletto, D.N. Anstett, J. Anstett, A. Bucharova, M.S. Comerford, S. David, M. Falahati-Anbaran, W. Godsoe, C. González-Lagos, P.E. Gundel, G.R. Hood, C. Lampei, C. Lara, A. Lázaro-Lobo, D.S. Leandro, T.J.S. Merritt, N. Mitchell, M.M. Bazargani, A. Moles, M. Murúa, J. Paule, V. Pfeiffer, J.A.M. Raeymaekers, D.J. Rennison, R.S. Rios, J.K. Rowntree, A.C. Schneider, K.S. Whitney, Í. Tamburrino, A. VanWallendael, P.Y. Kim, R.W. Ness, M.T. J. Johnson, K.A. Hodgins & N.J. Kooyers. 2025. Haploblocks contribute to parallel climate adaptation following global invasion of a cosmopolitan plant. Nature Ecology & Evolution 9: 1441-1455.
Kooyers, N.J., J.T. Anderson, A.L. Angert, M.L. Avolio, D.R. Campbell, M. Exposito-Alonso, T.E. Juenger, D.A. Moeller, J.D. Napier, and S.N. Sheth. 2025. Responses to climate change – insights and limitations from herbaceous plant model species. New Phytologist, 248: 461-493.
Hendrickson B. T., C.Stamps, C.M. Patterson, H.Strickland, M.Foster, L.J. Albano, A.Y. Kim, P.Y. Kim, N.J. Kooyers. 2025. Evolution of drought resistance strategies following the introduction of white clover (Trifolium repens L.). Annals of Botany, 135(7): mcaf037.
Mendez‐Reneau J.I., J.L. Richards, J. Hobbie, E. Bollich, N.J. Kooyers, E.M. Sigel. 2024. Lineage diversification and rampant hybridization among subspecies explain taxonomic confusion in the endemic Hawaiian fern Polypodium pellucidum. American Journal of Botany, 111(9): e16379.
Cocciardi J.M., A.M. Hoffman, D.F. Alvarado-Serrano, J. Anderson, M. Blumstein, E.L. Boehm, L.G. Bolin, I.T. Borokini, G.S. Bradburd, H.A. Branch,L.A. Brudvig, Y. Chen, S.L. Collins, D.L. Des Marais, D. Gamba, N.P. Hanan, M.M. Howard, J. Jaros, T.E. Juenger, N.J. Kooyers, E.J. Kottler, J.A. Lau, M. Menon, D.A. Moeller, T.J. Mozdzer, S.N. Sheth, M. Smith, K. Toll, M.C. Ungerer, M.L. Vahsen, S.M. Wadgymar, A. Waananen, K.D. Whitney, M.L. Avolio. 2024. The value of long-term ecological research for evolutionary insights. Nature Ecology & Evolution: 8, 1584–1592.
Santangelo J.S., P. Battlay, B.T. Hendrickson, W.H. Kuo, K.M. Olsen, N.J. Kooyers, M.T.J. Johnson, K.A. Hodgins, R.W. Ness. 2023. Haplotype-resolved, chromosome-level assembly of white clover (Trifolium repens L., Fabaceae). Genome Biology and Evolution. 15(8): 1-8
McDonald, L.M., A. Scharnagl, A.K. Turcu, C.M. Patterson, N.J. Kooyers. 2023. Demographic consequences of an extreme heatwave are mitigated by spatial heterogeneity in an annual monkeyflower. Ecology and Evolution 13: e10397
FitzPatrick, J.A., B.I. Doucet, S.D. Holt, C.M. Patterson, N.J. Kooyers. 2023. Unique drought resistance strategies occur among monkeyflower populations spanning an aridity gradient. American Journal of Botany, 110(8):e16207. Featured: AJB Highlights August 2023
Holt Jr, S.D., E.M. Sigel, B.L. Sutherland, P.B. Schwartsburd, J.B. Beck. 2023. What is Salvinia molesta (Salviniaceae)? Determining the maternal progenitor and genetic diversity of the clonal invasive fern giant salvinia. Biological Invasions 25(7): 2131-2141
Mendez-Reneau, J., J.G. Burleigh, E.M. Sigel. 2023. Target capture methods offer insight into the evolution of rapidly diverged taxa and resolve allopolyploid homeologs in the fern genus Polypodium ss. Systematic Botany.
Scharnagl, A., M.A. Genung, L.M. Holeski, N.J. Kooyers. 2023. Life history and chemical defense interact to drive patterns of adaptation in an annual monkeyflower. Evolution 77(2): 370-383
Innes, S.G., J.S. Santangelo, N.J. Kooyers, K.M. Olsen, M.T.J. Johnson. 2022. Evolution in response to climate in the native and introduced ranges of a globally distributed plant. Evolution 76(7): 1495-1511
Colicchio, J.M, L.N. Hamm, H.E. Verdonk, N.J. Kooyers, B.K. Blackman. 2021. Adaptive and non-adaptive causes of heterogeneity in genetic differentiation across the Mimulus guttatus. Molecular Ecology http://doi.org/10.1111/mec.16087
Sutherland B.L., C.F. Barrett, J.B. Beck, M. Latvis, M.R. McKain, E.M. Sigel, N.J. Kooyers. 2021. Botany is the Root and the Future of Invasion Biology. American Journal of Botany 108(4): 519-552
Paudel S., K. Bagale, S. Patel, N.J. Kooyers, R. Kulkarni. 2021. Human urine alters methicillin-resistant Staphylococcus aureus virulence and transcriptome. Applied and Environmental Microbiology. 87(16): e00744-21
Holeski, L.M., K. Keefover-Ring, J.M. Sobel, N.J. Kooyers. 2021. Evolutionary history and ecology shape the diversity and abundance of phytochemical arsenals across monkeyflowers. Journal of Evolutionary Biology 34(3): 571-583
Kooyers, N. J., K.A. Morioka, J.M. Colicchio, K.S. Clark, A. Donofrio, S.K. Estill, C.R. Pascualy, I.C. Anderson, M. Hagler, C. Cho, B.K. Blackman. 2021. Population responses to a historic drought across the range of the common monkeyflower (Mimulus guttatus). American Journal of Botany 108(2):1-13.
Kooyers, N.J., A. Donofrio, B.K. Blackman, L.M Holeski. 2020. The genetic architecture of plant defense tradeoffs in a common monkeyflower. Journal of Heredity 111(4):333-345 (Cover Article)
Kooyers, N.J. 2019. Are drought resistance strategies associated with life history strategy? Annals of Botany 124(1):vi-viii. Link
Lowry, D.B., J.M. Sobel, A.L. Angert, T.L Ashman, R.L. Baker, B.K. Blackman, Y. Brandvain, K.J.R.P. Byers, A.M. Cooley, J.M. Coughlan, M.R. Dudash, C.B. Fenster, K.G. Ferris, L. Fishman, J. Friedman, D.L. Grossenbacher, L.M. Holeski, C.T. Ivey, K.M. Kay, V.A. Koelling, N.J. Kooyers, M. Vallejo-Marín, C.J. Murren, M.L. Peterson, J.R. Puzey, M.C. Rotter, J.R. Seemann, J.P. Sexton , S.N. Sheth, M.A. Streisfeld, A.L. Sweigart, A.D. Twyford, J.H. Willis, C.A. Wu, Y.W Yuan. 2019. The case for the continued use of the genus name Mimulus for all monkeyflowers. Taxon 68(4): 617-623
Kooyers, N.J., J.M Colicchio, A.B. Greenlee, E. Patterson, N.T. Handloser, B.K. Blackman. 2019. Lagging adaptation to climate change supersedes local adaptation to herbivory in an annual monkeyflower. The American Naturalist. 194(4): 541-557
Kooyers, N.J., B.H. Bakken, M.C. Ungerer,K.M. Olsen. 2018. Freeze-induced cyanide toxicity does not maintain the cyanogenesis polymorphism in white clover (Trifolium repens). American Journal of Botany 105(7): 1-8 (Cover Article)
Kooyers, N.J., B.K. Blackman, L.M Holeski. 2017. Optimal defense theory explains deviations from latitudinal herbivory defense hypothesis. Ecology 98(4): 1036-1048
Kooyers, N.J., B. James, B.K. Blackman. 2017. Competition drives trait evolution and character displacement between Mimulus species along an environmental gradient. Evolution 71(5): 1425-1427
Kooyers, N.J. 2015. The evolution of drought escape and avoidance in natural herbaceous populations. Plant Science 234: 155-162
Kooyers, N.J., A.B. Greenlee, J.M. Colicchio, M. Oh, B.K. Blackman. 2014. Replicate altitudinal clines reveal evolutionary flexibility underlies adaptation to drought stress in annual Mimulus guttatus. New Phytologist. 206: 152-165
Kooyers, N.J., K.M. Olsen. 2014. Adaptive cyanogenesis clines in introduced regions evolve through geographical sorting of previously existing gene deletions. Journal of Evolutionary Biology: 27:2554-2558
Kooyers, N.J., L.R. Gage, A. Al-Lozi, K.M. Olsen. 2014. Aridity shapes cyanogenesis cline evolution in white clover (Trifolium repens L.). Molecular Ecology 23:1053-1070
Olsen, K.M., N .J. Kooyers, L. Small.2014 Adaptive gains through repeated gene loss: Parallel evolution of cyanogenesis polymorphisms in the genus Trifolium. Philosophical Transactions of the Royal Society B: Biological Sciences 369: 20130347.
Kooyers, N.J., K.M. Olsen. 2013. Searching for the Bull’s-eye: Targets of selection on cyanogenesis in white clover (Trifolium repens L.) vary between geographically disparate clinal replicates. Heredity. 11:495-504
Olsen, K.M., N.J. Kooyers, L. Small. 2012. Recurrent gene deletions and the evolution of adaptive cyanogenesis polymorphisms in white clover (Trifolium repens L.). Molecular Ecology 22: 724-738
Kooyers, N.J., K.M. Olsen. 2012. Rapid evolution of an adaptive cyanogenesis cline in introduced North American white clover (Trifolium repens L.) Molecular Ecology 21: 2455-2468
Dobritsa, A.A. , A. Geanconteri, J. Shrestha, A. Carlson, N.J. Kooyers*, D. Coerper, E. Urbanczyk, B.J. Bench, L.W. Sumner, R. Swanson, D. Preuss. 2011. A large-scale genetic screen in Arabidopsis thaliana to identify genes involved in pollen exine production. Plant Physiology 157: 947-970.
Gaspard, L.F., E. Harb, S.J. Holt Jr., C.M. Patterson, N.J. Kooyers. Accepted. Interactions between maternal provisioning and natural selection on seed mass fluctuate across heat waves. American Journal of Botany
Albano, L.J., C.M. Patterson, E.J. Kidder, N.J. Kooyers, M.T.J. Johnson. 2026. A field experiment assessing the roles of drought, herbivory, and local climate on cyanogenesis cline formation and local adaptation in Trifolium repens. Journal of Evolutionary Biology 39(4): 535-551
Albano, L.J., C.C. Bastias, A. Estarague, B.T. Hendrickson, S.G. Innes, N. King, C.M. Patterson, A. Tudoran, F. Vasseur, A. Puentes, C. Violle, N.J. Kooyers, M.T.J. Johnson. 2026. A transcontinental experiment elucidates (mal)adaptation of a cosmopolitan plant to climate in space and time. Ecological Monographs 96(1): e70043.
Innes, S.I., L.M. Holeski, N.J. Kooyers. 2025. Ontogenetic and geographic phytochemical variation in Mimulus moschatus, a perennial monkeyflower. Journal of Chemical Ecology. 51 (106): 1-12.
Kooyers, N. J., M.A. Genung, S.G. Innes, A.Turcu, D.M. Hinrichs, B.J. LeBlanc, and C.M. Patterson. 2025. Heat waves decrease fitness and alter maternal provisioning in natural populations of Mimulus guttatus. American Journal of Botany 112(8): e70087.
Battlay P., B.T. Hendrickson, J.I. Mendez-Reneau, J.S. Santangelo, L.J. Albano, J. Wilson, A.E. Caizergues, N. King, A. Puentes, A. Tudoran, C. Violle, F. Vasseur, C.M. Patterson, M. Foster, C. Stamps, S.G. Innes, R. Allio, F. Angeoletto, D.N. Anstett, J. Anstett, A. Bucharova, M.S. Comerford, S. David, M. Falahati-Anbaran, W. Godsoe, C. González-Lagos, P.E. Gundel, G.R. Hood, C. Lampei, C. Lara, A. Lázaro-Lobo, D.S. Leandro, T.J.S. Merritt, N. Mitchell, M.M. Bazargani, A. Moles, M. Murúa, J. Paule, V. Pfeiffer, J.A.M. Raeymaekers, D.J. Rennison, R.S. Rios, J.K. Rowntree, A.C. Schneider, K.S. Whitney, Í. Tamburrino, A. VanWallendael, P.Y. Kim, R.W. Ness, M.T. J. Johnson, K.A. Hodgins & N.J. Kooyers. 2025. Haploblocks contribute to parallel climate adaptation following global invasion of a cosmopolitan plant. Nature Ecology & Evolution 9: 1441-1455.
Kooyers, N.J., J.T. Anderson, A.L. Angert, M.L. Avolio, D.R. Campbell, M. Exposito-Alonso, T.E. Juenger, D.A. Moeller, J.D. Napier, and S.N. Sheth. 2025. Responses to climate change – insights and limitations from herbaceous plant model species. New Phytologist, 248: 461-493.
Hendrickson B. T., C.Stamps, C.M. Patterson, H.Strickland, M.Foster, L.J. Albano, A.Y. Kim, P.Y. Kim, N.J. Kooyers. 2025. Evolution of drought resistance strategies following the introduction of white clover (Trifolium repens L.). Annals of Botany, 135(7): mcaf037.
Mendez‐Reneau J.I., J.L. Richards, J. Hobbie, E. Bollich, N.J. Kooyers, E.M. Sigel. 2024. Lineage diversification and rampant hybridization among subspecies explain taxonomic confusion in the endemic Hawaiian fern Polypodium pellucidum. American Journal of Botany, 111(9): e16379.
Cocciardi J.M., A.M. Hoffman, D.F. Alvarado-Serrano, J. Anderson, M. Blumstein, E.L. Boehm, L.G. Bolin, I.T. Borokini, G.S. Bradburd, H.A. Branch,L.A. Brudvig, Y. Chen, S.L. Collins, D.L. Des Marais, D. Gamba, N.P. Hanan, M.M. Howard, J. Jaros, T.E. Juenger, N.J. Kooyers, E.J. Kottler, J.A. Lau, M. Menon, D.A. Moeller, T.J. Mozdzer, S.N. Sheth, M. Smith, K. Toll, M.C. Ungerer, M.L. Vahsen, S.M. Wadgymar, A. Waananen, K.D. Whitney, M.L. Avolio. 2024. The value of long-term ecological research for evolutionary insights. Nature Ecology & Evolution: 8, 1584–1592.
Santangelo J.S., P. Battlay, B.T. Hendrickson, W.H. Kuo, K.M. Olsen, N.J. Kooyers, M.T.J. Johnson, K.A. Hodgins, R.W. Ness. 2023. Haplotype-resolved, chromosome-level assembly of white clover (Trifolium repens L., Fabaceae). Genome Biology and Evolution. 15(8): 1-8
McDonald, L.M., A. Scharnagl, A.K. Turcu, C.M. Patterson, N.J. Kooyers. 2023. Demographic consequences of an extreme heatwave are mitigated by spatial heterogeneity in an annual monkeyflower. Ecology and Evolution 13: e10397
FitzPatrick, J.A., B.I. Doucet, S.D. Holt, C.M. Patterson, N.J. Kooyers. 2023. Unique drought resistance strategies occur among monkeyflower populations spanning an aridity gradient. American Journal of Botany, 110(8):e16207. Featured: AJB Highlights August 2023
Holt Jr, S.D., E.M. Sigel, B.L. Sutherland, P.B. Schwartsburd, J.B. Beck. 2023. What is Salvinia molesta (Salviniaceae)? Determining the maternal progenitor and genetic diversity of the clonal invasive fern giant salvinia. Biological Invasions 25(7): 2131-2141
Mendez-Reneau, J., J.G. Burleigh, E.M. Sigel. 2023. Target capture methods offer insight into the evolution of rapidly diverged taxa and resolve allopolyploid homeologs in the fern genus Polypodium ss. Systematic Botany.
Scharnagl, A., M.A. Genung, L.M. Holeski, N.J. Kooyers. 2023. Life history and chemical defense interact to drive patterns of adaptation in an annual monkeyflower. Evolution 77(2): 370-383
Innes, S.G., J.S. Santangelo, N.J. Kooyers, K.M. Olsen, M.T.J. Johnson. 2022. Evolution in response to climate in the native and introduced ranges of a globally distributed plant. Evolution 76(7): 1495-1511
Colicchio, J.M, L.N. Hamm, H.E. Verdonk, N.J. Kooyers, B.K. Blackman. 2021. Adaptive and non-adaptive causes of heterogeneity in genetic differentiation across the Mimulus guttatus. Molecular Ecology http://doi.org/10.1111/mec.16087
Sutherland B.L., C.F. Barrett, J.B. Beck, M. Latvis, M.R. McKain, E.M. Sigel, N.J. Kooyers. 2021. Botany is the Root and the Future of Invasion Biology. American Journal of Botany 108(4): 519-552
Paudel S., K. Bagale, S. Patel, N.J. Kooyers, R. Kulkarni. 2021. Human urine alters methicillin-resistant Staphylococcus aureus virulence and transcriptome. Applied and Environmental Microbiology. 87(16): e00744-21
Holeski, L.M., K. Keefover-Ring, J.M. Sobel, N.J. Kooyers. 2021. Evolutionary history and ecology shape the diversity and abundance of phytochemical arsenals across monkeyflowers. Journal of Evolutionary Biology 34(3): 571-583
Kooyers, N. J., K.A. Morioka, J.M. Colicchio, K.S. Clark, A. Donofrio, S.K. Estill, C.R. Pascualy, I.C. Anderson, M. Hagler, C. Cho, B.K. Blackman. 2021. Population responses to a historic drought across the range of the common monkeyflower (Mimulus guttatus). American Journal of Botany 108(2):1-13.
Kooyers, N.J., A. Donofrio, B.K. Blackman, L.M Holeski. 2020. The genetic architecture of plant defense tradeoffs in a common monkeyflower. Journal of Heredity 111(4):333-345 (Cover Article)
Kooyers, N.J. 2019. Are drought resistance strategies associated with life history strategy? Annals of Botany 124(1):vi-viii. Link
Lowry, D.B., J.M. Sobel, A.L. Angert, T.L Ashman, R.L. Baker, B.K. Blackman, Y. Brandvain, K.J.R.P. Byers, A.M. Cooley, J.M. Coughlan, M.R. Dudash, C.B. Fenster, K.G. Ferris, L. Fishman, J. Friedman, D.L. Grossenbacher, L.M. Holeski, C.T. Ivey, K.M. Kay, V.A. Koelling, N.J. Kooyers, M. Vallejo-Marín, C.J. Murren, M.L. Peterson, J.R. Puzey, M.C. Rotter, J.R. Seemann, J.P. Sexton , S.N. Sheth, M.A. Streisfeld, A.L. Sweigart, A.D. Twyford, J.H. Willis, C.A. Wu, Y.W Yuan. 2019. The case for the continued use of the genus name Mimulus for all monkeyflowers. Taxon 68(4): 617-623
Kooyers, N.J., J.M Colicchio, A.B. Greenlee, E. Patterson, N.T. Handloser, B.K. Blackman. 2019. Lagging adaptation to climate change supersedes local adaptation to herbivory in an annual monkeyflower. The American Naturalist. 194(4): 541-557
Kooyers, N.J., B.H. Bakken, M.C. Ungerer,K.M. Olsen. 2018. Freeze-induced cyanide toxicity does not maintain the cyanogenesis polymorphism in white clover (Trifolium repens). American Journal of Botany 105(7): 1-8 (Cover Article)
Kooyers, N.J., B.K. Blackman, L.M Holeski. 2017. Optimal defense theory explains deviations from latitudinal herbivory defense hypothesis. Ecology 98(4): 1036-1048
Kooyers, N.J., B. James, B.K. Blackman. 2017. Competition drives trait evolution and character displacement between Mimulus species along an environmental gradient. Evolution 71(5): 1425-1427
- Featured: K.E. Eisen. 2017. Digest: Trait variation in Mimulus provides new evidence for the joint action of ecological sorting and character displacement. Evolution 71(5): 1425-1427
Kooyers, N.J. 2015. The evolution of drought escape and avoidance in natural herbaceous populations. Plant Science 234: 155-162
Kooyers, N.J., A.B. Greenlee, J.M. Colicchio, M. Oh, B.K. Blackman. 2014. Replicate altitudinal clines reveal evolutionary flexibility underlies adaptation to drought stress in annual Mimulus guttatus. New Phytologist. 206: 152-165
Kooyers, N.J., K.M. Olsen. 2014. Adaptive cyanogenesis clines in introduced regions evolve through geographical sorting of previously existing gene deletions. Journal of Evolutionary Biology: 27:2554-2558
Kooyers, N.J., L.R. Gage, A. Al-Lozi, K.M. Olsen. 2014. Aridity shapes cyanogenesis cline evolution in white clover (Trifolium repens L.). Molecular Ecology 23:1053-1070
Olsen, K.M., N .J. Kooyers, L. Small.2014 Adaptive gains through repeated gene loss: Parallel evolution of cyanogenesis polymorphisms in the genus Trifolium. Philosophical Transactions of the Royal Society B: Biological Sciences 369: 20130347.
Kooyers, N.J., K.M. Olsen. 2013. Searching for the Bull’s-eye: Targets of selection on cyanogenesis in white clover (Trifolium repens L.) vary between geographically disparate clinal replicates. Heredity. 11:495-504
Olsen, K.M., N.J. Kooyers, L. Small. 2012. Recurrent gene deletions and the evolution of adaptive cyanogenesis polymorphisms in white clover (Trifolium repens L.). Molecular Ecology 22: 724-738
Kooyers, N.J., K.M. Olsen. 2012. Rapid evolution of an adaptive cyanogenesis cline in introduced North American white clover (Trifolium repens L.) Molecular Ecology 21: 2455-2468
- Featured: Molecular Ecology Perspective, Olson, M.S., N. Levsen (2012) Classic clover cline clues. Molecular Ecology 21: 2315-2317
Dobritsa, A.A. , A. Geanconteri, J. Shrestha, A. Carlson, N.J. Kooyers*, D. Coerper, E. Urbanczyk, B.J. Bench, L.W. Sumner, R. Swanson, D. Preuss. 2011. A large-scale genetic screen in Arabidopsis thaliana to identify genes involved in pollen exine production. Plant Physiology 157: 947-970.