Nyree Zerega, Ph.D.

Nyree Zerega
Director, Graduate Program in Plant Biology and Conservation; Herbarium Director; Conservation Scientist; Professor of Instruction at Northwestern University
(847) 835-6916
Teaching and Research Affiliations:
Research Interests:
  • Evolutionary processes in plants, especially as they apply to crops and their wild relatives
  • Origins, diversity, and conservation of underutilized crop plants, especially in the genus Artocarpus 
  • Plant systematics and biogeography
  • Pollination biology
  • Perennial crops (including breadfruit, jackfruit, and Kernza) and sustainable agriculture

Lab webpage


Inspired by the amazing diversity and utility of plants, my research uses phylogenomic, collections-based, and experimental field approaches to explore the evolution and genetic diversity of underutilized crops and their wild relatives, pollination, biogeography, and taxonomic revisionary work. I am also interested in applying findings to conservation of plant genomic resources and more sustainable agriculture. 

Selected Publications:

Linkd to Google Scholar for updates

Gardner, E.M., S. Bruun-Lund, M.A. Niissalo, B,Chantarasuwan, W.L. Clement, C. Geri, R.D. Harrison, A.L. Hipp, M. Holvoet, G. Khew, F, Kjellberg, S. Liao, L.C. Pederneiras, Y-Q. Peng, J.T. Pereira, Q. Phillips, A.S.A. Puad, J-Y. Rasplus, J. Sang, S. Schou, E. Velautham, G.D. Weiblen, N.J.C. Zerega, Q. Zhang, Z. Zhang. C. Baraloto, N. Rønsted. 2023. Echoes of ancient introgression punctuate stable genomic lineages in the evolution of figs. Proceedings of National Academy of Sciences

Audi, L., G. Shallow, E. Robertson, D. Bobo, D. Ragone, E.M. Gardner, B. Dussoruth, J. Wajer, and N.J.C. Zerega. 2023 Linking breadfruit cultivar names across the globe connects histories after 230 years of separation. Current Biology

Chamberlain, L.A, T. Aguayo, N.J.C. Zerega, R. Dybzinksi, and L. Egerton-Warburton. 2022. Rapid improvement in soil health following the conversion of abandoned farm fields to annual or perennial agroecosystems. Frontiers in Sustainable Food Systems. 6:1010298. 

Chuang C-R, Hsieh C-L, Chang C-S, Wang C-M, Tandang DN, Gardner EM, et al. 2022. Amis Pacilo and Yami Cipoho are not the same as the Pacific breadfruit starch crop—Target enrichment phylogenomics of a long-misidentified Artocarpusspecies sheds light on the northward Austronesian migration from the Philippines to Taiwan. PLoS ONE 17(9): e0272680.

Yang L, Zerega NJC, Montgomery A, Horton DE. 2022. Potential of breadfruit cultivation to contribute to climate-resilient low latitude food systems. PLOS Clim 1(8): e0000062.

Gardner, E.M.,S.A. Puad, J.T. Pereira, L. Pokorny, and N.J.C. Zerega. 2022. Engagement with indigenous knowledge improves our understanding of biodiversity and promotes the conservation of both. Current Biology.

Iwanycki Ahlstrand Natalie, et al, N.J.C. Zerega, Rønsted, N. 2022. Travel tales of a worldwide weed: genomic signature of Plantago major reveal distinct genotypic groups with links to colonial trade routes. Frontiers in Plant Science 13

Gardner, E.M., T. Jimbo, and N.J.C. Zerega. 2022. Artocarpus buyangensis, a new species from New Guinea. Systematic Botany 47(2): 452-456

Gardner, E.M., M. Garner, R. Cowan, S. Dodsworth, N. Epitawalage, O. Maurin, D. Arifiani, S. Sahromi, W.J. Baker, F. Forest, N.J.C. Zerega, A.K. Monro, A.L. Hipp. 2021. Repeated parallel losses of inflexed stamens in Moraceae: phylogenomics and generic revision of the tribe Moreae and the reinstatement of the tribe Olmedieae (Moraceae). Taxon.

Lindsay, D.J. et al., and N.J.C. Zerega, R.-L. Zhu, K.B.H. Er. 2022. Flora of Singapore: Checklist and bibliography. The Gardens’ Bulletin Singapore 74: 2383-5812.’-bulletin-singapore/listing-of-publications

Marion Suiseeya, K. M., G. O’Connell, E. Leoso, S.B.N. DeFoe, A. Anderson, M. Bang, P. Beckman, A. Boyer, J. Dunn, J. Gilbert, J. Hester, D.E. Horton, D.B. Jennings, P. Kebec, N. Loeb, P. Loew, W.M. Miller, K. Moffit, A.I. Packman, M.W. Price, B. Redbird, J. Rogers, R. Sankaran, J. Schwoch, P. Silas, W. Twardowski, N.J.C. Zerega. 2022. Waking from Paralysis – Revitalizing Conceptions of Climate Knowledge and Justice for More Effective Climate Action. Annals of the American Academy of Political and Social Science

Gardner, E.M. and N.J.C. Zerega. 2021. Taxonomic updates to Artocarpus subgenus Artocarpus (Moraceae)and allied taxa with a particular focus on the species native to Singapore. Gardens’ Bulletin Singapore 73(2): 309-374.

Gardner, E.M., M. Garner, R. Cowan, S. Dodsworth, N. Epitawalage, O. Maurin, D. Arifiani, S. Sahromi, W.J. Baker, F. Forest, N.J.C. Zerega, A.K. Monro, A.L. Hipp. 2021. Repeated parallel losses of inflexed stamens in Moraceae: phylogenomics and generic revision of the tribe Moreae and the reinstatement of the tribe Olmedieae (Moraceae). Taxon.

Gardner, E.M., L. Audi, Q. Zhang, H. Sauquet, A.K. Monro, and N.J.C. Zerega. 2021Phylogenomics of Brosimum Sw. (Moraceae) and allied genera, including a revised subgeneric system. Taxon. 

Gardner, E.M., D. Arifiani, and N.J.C. Zerega. 2021. Artocarpus bergii (Moraceae), a new species in the breadfruit clade from the Moluccas. Systematic Botany 46(1) (2021).  

Gardner, E.M. and N.J.C. Zerega. 2020. Taxonomic updates to Artocarpus subgenus Pseudojaca (Moraceae), with a particular focus on the taxa in Singapore. Gardens’ Bulletin Singapore 72(2): 173-213. doi: 10.26492/gbs72(2).2020-06

Gardner, E.M., N.J.C. Zerega, A.K. Monro. 2020. (2757) Proposal to reject the name Antiarideae (Moraceae). Taxon 69(4):827-828.  

Gardner, E.M., M.G. Johnson, J.T. Pereira, A.S.A. Puad, D. Arifiani, Sahromi, N.J. Wickett, and N.J.C. Zerega. 2021. Paralogs and off-target sequences improve phylogenetic resolution in a densely-sampled study of the breadfruit genus (Artocarpus, Moraceae). Systematic Biology syaa073,

Gardner, E.M. and N.J.C. Zerega. 2020. Proposal to reject the name Radermachia rotunda (Artocarpus rotundus) (Moraceae) (2788). Taxon 69(4): 827-828.

Gardner, E.M., A. Chaveerach, R. Sudmoon, N.J.C. Zerega. 2020. Two new species of Artocarpus (Moraceae) from Thailand and VietnamPhytotaxa Vol. 453, No. 3.

Dickinson, L., H. Noble, E.M. Gardner, A.S.A. Puad, Wan Zakaria, W. and N.J.C. Zerega. 2020. Genetic diversity structure and conservation of critically endangered, Artocarpus annulatus, a crop wild relative of jackfruit. PeerJ. 10.7717/peerj.9897

Zerega, N.J.C., Gardner, E., Wan Zakaria, W. & Ahmad Puad, A.S. 2019. Artocarpus annulatus. The IUCN Red List of Threatened Species2019: e.T86528602A86528612.

Zhang, Q., E.M Gardner, N.J.C. Zerega, and H. Sauquet. 2019. Long-distance dispersal shaped the diversity of tribe Dorstenieae (Moraceae). Available at BioRxiv

Witherup, C.E., M.I. Zuberi, S. Hossain, and N.J.C. Zerega. 2019. Genetic Diversity of Bangladeshi Jackfruit (Artocarpus heterophyllus) over Time and Across Seedling Sources. Journal of Economic Botany

Sahu, S.k., M. Liu, Yssel, R. Kariba, S. Jiang, B. Song, S. Muthemba, P.S. Hendre, R. Jamnadass, S.M. Kao, N.J.C. Zerega, X. Xu, H. Yang, A. Van Deynze, Y. Van de Peer, X. Liu and H. Liu. 2020. Draft Genomes of two Artocarpus plants, Jackfruit (A. heterophyllus) and Breadfruit (A. altilis). Genes 11(1). pii: E27.

Witherup, C.E., M.I. Zuberi, S. Hossain, and N.J.C. Zerega. 2019. Genetic Diversity of Bangladeshi Jackfruit (Artocarpus heterophyllus) over Time and Across Seedling Sources. Journal of Economic Botany

Zerega, N.J.C andE.M. Gardner. Delimitation of the new tribe Parartocarpeae (Moraceae) is supported by a 333-gene phylogeny and resolves tribal level Moraceae taxonomy. 2019. Phytotaxa 388 (4): 253–265.

Lincoln, N.K., D. Ragone, N.J.C. Zerega, L. Nkrumah-Roberts, M. Merlin, and A.M. Jones. 2018. Grow Us Our Daily Bread: A Review of Breadfruit Cultivation in Traditional and Contemporary Systems. Horticultural Review 46:299-384.

Laricchia, K.M., M. Johnson, N.J.C. Zerega, and N.J. Wickett. 2018. A transcriptome screen for positive selection in domesticated breadfruit and its wild relatives (Artocarpus spp.). American Journal of Botany.  

Wang, M., E.M. Gardner, A.R. Milan, Salma, C.M. Yee, R. Chung, L.S. Leong, and N.J.C. Zerega. 2018. Origin and diversity of an underutilized fruit tree crop, Artocarpus integer(Moraceae). American Journal of Botany.

Gardner, E.M. R. Gagne, P.E. Kendra, W.S. Montgomery, R.A. Raguso, T.T. McNeil and N.J.C. Zerega. A flower in fruit’s clothing: Pollination of jackfruit (Artocarpus heterophyllus, Moraceae) by a new species of gall midge,Clinodiplosis ultracrepidatasp. nov. (Diptera: Cecidomyiidae). 2018. International Journal of Plant Sciences.

Kates, H.R., M.G. Johnson, E.M. Gardner, N.J.C. Zerega, and N.J. Wickett. 2018. Allele phasing has minimal impact on phylogenetic reconstruction from targeted nuclear gene sequences in a case study of Artocarpus(Moraceae). American Journal of Botany

Briscoe, L.R.E., N.J.C. Zerega, H.T. Lumbsch, M. Stech, E. Kraichak, M.J. von Konrat, J.J. Engel, and N. Wickett. 2017. Molecular, morphological, and biogeographic perspectives on the classification of Acrobolboideae (Acrobolbaceae, Marchantiophyta). Phytotaxa319 (1): 056–070.

Gardner, E.M., P. Sarraf, and N.J.C. Zerega. Phylogeny and biogeography of Maclura (Moraceae) and the origin of an anachronistic fruit. 2017. Molecular Phylogenetics and Evolution.

Williams, E.W., E.M. Gardner, R. Harris III, A. Chaveerach, J.T. Pereira, and N.J.C. Zerega. Out of Borneo: Biogeography, phylogeny, and divergence date estimates of Artocarpus (Moraceae). Accepted pending minor revisions. Annals of Botany.

Zerega, N.J.C. and D. Ragone. In press. Toward a global view of breadfruit genetic diversity. Proceedings from the International Breadfruit Conference. Tropical Agriculture.

Gardner, E.M., M.G. Johnson, D. Ragone, N.J. Wickett, & N.J.C. Zerega. 2016. Low-coverage, whole-genome sequencing of Artocarpus camansi(Moraceae) for phylogenetic marker development and gene discovery. Applications in Plant Sciences4(7), apps.1600017.

Johnson, M.G., E.M. Gardner, Y. Liu, R. Medina, B. Goffinet, A.J., N.J.C. Zerega, and N.J. Wickett. 2016. HybPiper: Extracting coding sequence and introns for phylogenetics from high-throughput sequencing reads using target enrichment.Applications in Plant Sciences4(7), apps.1600016.

Gardner, E.M., K.M. Laricchia, M. Murphy, D. Ragone, B.E. Scheffler, S. Simpson, E.W. Williams, and N.J.C. Zerega. 2015. Primer Note: Chloroplast microsatellite markers for Artocarpus (Moraceae) developed from transcriptome sequences. Applications in Plant Sciences 3(9):

Zerega, N.J.C., T. Wiesner-Hanks, D. Ragone, B. Irish, B. Scheffler, S. Simpson, and F. Zee. 2015. Diversity of the breadfruit complex (Artocarpus, Moraceae): Genetic characterization of critical germplasm. Tree Genetics and Genomes.  

Zerega, N.J.C., G. Ravikanth, E.M. Gardner, K. Laricchia, T. Melhem, M. Wang, C. E. Witherup, S. Setty, M.I. Zuberi, M.R. Jagdish, and D. Ragone. 2015. New genomic resources to understand genetic diversity and promote crop improvement of jackfruit (Artocarpus heterophyllus), breadfruit (A. altilis), and other under utilized Artocarpus crops. Procedings of the International Symposium on Jackfruit and Breadfruit of the Tropics.

Zerega, N.J.C. 2014. Origins and Development of Breadfruit, in C. Smith (ed.). Encyclopedia of Global Archaeology. P. 234. Springer, New York, New York.

Witherup, C, D. Ragone, T. Wiesner-Hanks, B. Irish, B. Scheffler, S. Simpson, F. Zee, M. I. Zuberi, N.J.C. Zerega. 2013. "Development of microsatellite loci in Artocarpus altilis (Moraceae) and cross-amplification in congeneric species." Applications in Plant Science 1(7):

Misiewicz, T. and N.C. Zerega. 2012. "Phylogeny, Biogeography and Character Evolution of Dorstenia (Moraceae)." Edinburgh Journal of Botany 69(3): 413-440.

Zerega, N.J.C., M.N. Nur Supardi, and T.J. Motley. 2010. Phylogeny and recircumscription of Artocarpeae (Moraceae) with a focus on ArtocarpusSystematic Botany 35 (4): 766 - 783.

Khan, R., N.J.C. Zerega, S. Hossain, and M.I. Zuberi. 2010. Jackfruit (Artocarpus heterophyllus Lam.) Diversity in Bangladesh: Land Use and Artificial SelectionEconomic Botany 64(2): 124-136.

Rønsted, N., G.D. Weiblen, W.L. Clement, N.J.C. Zerega, and V. Savolainen. 2008. Reconstructing the phylogeny of figs (Ficus, Moraceae) to reveal the history of the fig pollination mutualism. Symbiosis 45:45-56.

Zerega, N.J.C., D. Ragone, and T.J. Motley. 2006. Genetic diversity and origins of domesticated breadfruit. In Darwin’s Harvest: New Approaches to the Origins, Evolution, and Conservation of Crops, ed. T.J. Motley, N.J.C. Zerega, and H.B. Cross. Columbia University Press, New York.

Motley, T.J., N.J.C. Zerega, and H.B. Cross, ed. 2006. Darwin’s Harvest: New Approaches to the Origins, Evolution, and Conservation of Crops. New York: Columbia University Press.

Zerega, N.J.C., W.L. Clement, S.L. Datwyler, and G.D. Weiblen. 2005. Biogeography and divergence times in the mulberry family (Moraceae)Molecular Phylogenetics and Evolution 37:402-416.

Zerega, N.J.C., D. Ragone, and T.J. Motley. 2005. Systematics and species limits of breadfruit (Artocarpus, Moraceae)Systematic Botany 30(3):603-615.

Zerega, N.J.C., L.A. Mound, and G.D. Weiblen. 2004. Pollination in the New Guinea endemic Antiaropsis decipiens (Moraceae) is mediated by a new species of thrips, Thrips antiaropsidis sp. nov. (Thysanoptera:Thripidae)International Journal of Plant Sciences 165(6):1,017-1,026.

Zerega, N.J.C., D. Ragone, and T.J. Motley. 2004. Complex origins of breadfruit: Implications for human migrations in OceaniaAmerican Journal of Botany 91(5):760-766.

Zerega, N.J.C. 2003. The Breadfruit TrailNatural History 112(10):46-51.

Zerega, N.J.C., S. Mori, C. Lindqvist, Q. Zheng, and T.J. Motley. 2002. Using amplified fragment length polymorphisms (AFLP) to identify black cohosh (Actaea racemosa)Economic Botany 56(2):154-164.

Bultman, T.L. and N.J. Conard. 1998. Effects of endophytic fungus, nutrient level, and plant damage on performance of Fall Armyworm (Lepidoptera: Noctuidae). Environmental Entomology 27(3):631-635.

Graduate Students:
Julie Phipps (Master's, expected graduation 2024)
Gretta Buttelmann (Master's, expected graduation 2024)
Evana James (PhD, expected graduation 2028)
Elise Neal (undergraduate, expected graduation 2026)
Samantha Turner-Rosa (Master's, 2024)
Lindsey Gohd (Master's, expected graduation 2021)
Leta Dickinson (Bachelor's degree 2019)
Lauren Audi (Master's 2018)
Elliot Gardner (Ph.D. 2017)
Theresa Melhem (Master's 2015)
Kristen Laricchia (Master's 2014)
Maria Wang (Master's 2014)
Paya Sarraf (Bachelor's degree 2014)
Tyr Wiesner-Hanks (Bachelor's degree 2013)
Colby Witherup (Master's degree 2013)
Lindsay Darling (Master's degree 2013)
Diane Huebner (Master's degree 2009)
Tracy Misiewicz (Master's degree 2009)
Evelyn Williams (Postdoctoral Research Associate, 2012-2014)

Zerega Lab Group
The Zerega Lab aims to understand evolutionary processes in plants, especially as they apply to underutilized crops and their wild relatives. The research takes both a basic and applied approach with the goals of advancing crop improvement, sustainable agriculture, and food security, as well as informing the conservation of plant genetic resources. 

Graduate Program in Plant Biology and Conservation
This program is a collaboration between Northwestern University and the Chicago Botanic Garden. The graduate program offers thesis and non-thesis master’s degree and Ph.D. programs and aims to foster an academic and research environment that allows students to gain experience, skills, and knowledge to become scholars, leaders, and practitioners in plant biology and conservation.

New Roots for Restoration

A collaborative NSF funded project advancing the understanding of diverse perennial systems in nature and agriculture