Plant Evolution
Professor & Curator, UC/JEPS
Lab Phone: 

Current Position

Professor, Departments of Plant and Microbial Biology & Integrative Biology

Curator of monocots, the University and Jepson Herbaria

Associate Editor, AoB PLANTS

Chief Editor, Frontiers for Young Minds, Biodiversity Section

Ph.D.  Plant Systematics and Evolution    New York University, 2004
B.A.   Biology    University of Delaware, 1993

The Evolution of Form and Function in Flowering Plants
Systematics and Evolution of the Spiral Gingers
My research with the “spiral gingers” involves utilizing a phylogenetic framework to investigate molecular evolution, floral development, and the historical biogeography of the pan-tropical family Costaceae. This family forms an easily recognizable clade within the Zingiberales, distinguished from other families in the order by its characteristic one-sided spiral phyllotaxy and its floral design, featuring 5 infertile stamens fused to form a petaloid labellum. Costaceae presently consists of about 95 species which until recently were placed in four genera: Costus, Tapeinochilos, Dimerocostus, and Monocostus. Recent systematic and developmental studies have led to a novel understanding of evolution within the family and have resulted in the description of three new genera (Chamaecostus, Cheilocostus and Paracostus) to accommodate the phylogenetic diversity of the Costaceae.

Costus lucanusianus, a monocot with a bee pollination syndrome.


The diverse floral morphology found in the family Costaceae makes this group an ideal model system for addressing a number of evolutionary questions. Research on this group revolves around four integrated components:
  • developing a robust phylogeny using molecular and morphological characters and revising the taxonomy of the lineage,
  • investigating floral pattern formation using taxa which encompass a range of morphological and phylogenetic diversity,
  • conducting comparative studies of floral development characters in order to explore the association between pollination syndrome, species diversity and rates of diversification, and
  • comparing the age of phylogenetic diversification with the geological record to interpret the biogeographic history of this group.


Costus pulverulentus, a monocot with a hummingbird pollination syndrome.


Evolution and Diversification of Monocots:
Evolution and Floral development in Zingiberales
My lab has several projects focusing on systematics and floral development in the order Zingiberales (tropical gingers). Programs looking at evolutionary patterns in the Heliconiaceae (Heliconia) and Zingiberaceae (Alpinioideae, Zingiberoideae) are underway. The results of these systematic studies are interpreted with respect to pollination biology, floral development, biogeography and molecular evolution at the ordinal level. Ongoing studies also include the evolution of floral development in the ginger order.
Evolution and Edaphic Variation in the Milla Complex
In collaboration with Dr. Victoria Sosa at INECOL, Xalapa, Mexico, we are investigating the systematic relationships of plants in the Milla complex of the monocot family Themidaceae (Asparagales). This includes genera Milla, Petronymphe, Jaimehintonia, Dandya and Bessera from Mexico as well as several groups from California. This project is funded by the UC MEXUS program.
A Milla from Mexico.
Evolution and Origin of California Alliums
With funding from the Heckard Fund, we are investigating the evolutionary position of California alliums (wild onions) within the worldwide genus (Allium). Preliminary results indicate that the California species may not be monophyletic and suggest an interesting history of adaptive radiation onto specialized serpentine soils.
Allium brevistylum from Utah and Colorado.
Phylogeography and Species Conservation
I am interested in biogeographic studies at the population-level, using molecular data to investigate distributional patterns within species and species complexes. Such studies can be combined with analyses of range reductions and genetic diversity among populations in order to provide information for informed conservation planning at the species level and to study the origins of crop plants to help direct genomic research of crop development. Current projects in this area include Pollination, Gene flow and Population genetics of the Mexican cycad genus Dioon.
Dioon purpusii in Oaxaca, Mexico.
Photo courtesy of the Palm & Cycad Societies of Florida, Inc.
web page (www.plantapalm.com).
Recent Publications

Thiago Andre**, Shayla Salzman+, Tania Wendt, and C.D. Specht. Speciation dynamics and biogeography of Neotropical spiral gingers (Costaceae). Molecular Phylogenetics and Evolution. doi:10.1016/j.ympev.2016.07.008

Chodon Sass, Will J.D. Iles, Craig Barrett, Selena Y. Smith, and C.D. Specht. 2015. Revisiting the Zingiberales: Using multiplexed exon capture to resolve ancient and recent phylogenetic splits in a charasmatic plant lineage.  PeerJ. 4-e1584.

Specht, C.D. The rise of evo-devo: the Pan American Society for Evolutionary Developmental Biology sets the stage. Am. J. Bot. 103: 1-4.

Kelsie Morioka+, Roxana Yockteng, Ana M.R. Almeida* and C.D. Specht. 2015. Loss of YABBY2-like gene expression may underlie the evolution of the laminar style in Canna and contribute to floral morphological diversity in the Zingiberales. Frontiers in Plant Science. http://dx.doi.org/10.3389/fpls.2015.01106

John C. Benedict, Selena Y. Smith, Margaret E. Collinson, Jana Leong-Škorničková, C. D. Specht, Julie L. Fife, Federica Marone, Xianghui Xiao, and Dilworth Y. Parkinson. 2015. Seed morphoanatomy and its utility in recognizing subfamilies and tribes of the Zingiberaceae. American Journal of Botany.

Moczek, A.P. et al. 2015.  The significance and scope of evolutionary developmental biology: a vision for the 21st century.  Evolution & Development. 17(13): 198-219. DOI:10.111/ede.12125.

Andre, Thiago, C.D. Specht, S. Salzman, Clarisse Palma-Silva, and Tania Wendt. 2015. Evolution of species diversity in the genus Chamaecostus (Costaceae): molecular phylogenetics and morphometric approaches. Phytotaxa 204(4): 265-276. doi: 10.11646/phytotaxa.204.4.3

Almeida, A.M.R., Yockteng, R., Otoni, W.C., and C.D. Specht. 2015. Positive selection on the K domain of the AGAMOUS protein in Zingiberales suggests a mechanism for the evolution of androecial morphology in the order. BMC EvoDevo.

A.M.R. Almeida, R. Yockteng and C.D. Specht. 2015. Evolution of petaloidy in the Zingiberales: an assessment of the relationship between untrastructure and gene expression patterns. Developmental Dynamics (special issue on Evolution & Morphological Diversity).

John C. Benedict, Selena Y. Smith, Margaret E. Collinson, Jana Leong-Škorničková, C. D. Specht, Julie L. Fife, Federica Marone, Xianghui Xiao, and Dilworth Y. Parkinson. 2015. Evolutionary significance of seed structure in Alpiniodeae (Zingiberaceae). Botanical Journal of the Linnean Society. 178 (3): 441-466. Doi: 10.111/boj.12257.

A.R. Deans et al. 2015. Finding our way through phenotypes. PloS Biology. DOI: 10.1371/journal.pbio.1002033.

Shayla Salzman+, Heather Driscoll, Tanya Renner*, Thiago Andre**, Stacy Shen*, and C. D. Specht. 2015. Spiraling into History: A molecular phylogeny and investigation of biogeographic origins of floral evolution for the genus Costus (Costaceae). Systematic Botany. 40(1): 104-115. DOI 10.1600/036364415X686404.

C.D. Specht & Dianella Howarth. 2015. Adaptation of floral form: an evodevo perspective. Tansley Review: New Phytologist. doi: 10.1111/nph.13198

Yockteng, R., Ana M.R. Almeida, Alma Pineyro-Nelson and C.D. Specht. 2014. Chapter 7: Adaptation of floral form: an evo-devo approach to study adaptive evolution in floral morphology. In Mechanisms in Plant Adaptation Roosa Laitinen (ed). Wiley Press.

Ana M.R. Almeida*, Roxana Yockteng, James C. Schnable**, Elena R. Alvarez-Buylla, Michael Freeling & C. D. Specht.  2014. Co-option of the polarity gene network shapes filament morphology in the angiosperms.  Nature: Scientific Reports, 4: 6194. doi:10.1038/srep06194.

Etelvina Gandara**, C.D. Specht and Victoria Sosa. 2014. Origin and diversification of the Milla clade (Brodiaeoideae, Asparagaceae): a neotropical group of six geophyte genera.  Molecular Phylogenetics and Evolution. Doi: 10.1016/j.ympev.2014.02.014

Craig F. Barrett, C. D. Specht, Jim Leebens-Mack, Dennis Wm. Stevenson, Wendy B. Zomlefer and Jerrold I. Davis. 2014. Resolving ancient radiations: can more data help resolve deep relationships among the tropical gingers (Zingiberales Griseb.). Annals of Botany, 113: 119–133.  doi:10.1093/aob/mct264

Eduardo Ruiz-Sanchez & C. D. Specht. 2014. Ecological speciation in Nolina parviflora (Asparagaceae): Lacking spatial connectivity along the Trans-Mexican Volcanic Belt. PloS ONE. 9(6):e98754. DOI:10.1371/journal.pone.0098754.

Roxana Yockteng, Ana M.R. Almeida*, Stephen Yee*, Thiago Andre**, Colin Hill+ and C. D. Specht.  2013. A method for extraction high quality RNA for next-generation sequencing and gene expression analyses.  Applications in Plant Sciences. 1(12). DOI: 10.3732/apps1300070.

Roxana Yockteng, Ana M.R. Almeida*, Kelsie Morioka+,Elena R. Alvarez-Buylla & C.D. Specht. 2013. Molecular evolution and patterns of duplication in the SEP/AGL6-like lineage of the Zingiberales: a proposed mechanism for floral diversification.  Molecular Biology and Evolution 30(11):2401–2422. DOI: 10.1093/molbev/mst137.

Tanya Renner & C.D. Specht. 2013. Inside the trap: gland morphologies, digestive enzymes, and the evolution of plant carnivory in the Caryophyllales.  Current Opinions in Plant Biology. DOI: 10.1016/j.pbi.2013.06.009.

Ana M.R. Almeida*, Andrew Brown+ & C.D. Specht.  2013. Tracking the development of the petaloid fertile stamen in Canna indica: Insights into the origin of androecial petaloidy in the Zingiberales. AoB PLANTS. DOI: 10.1093/aobpla/plt009.

Luke J. Harmon, J. Baumes, C. Hughes, J. Soberon, C.D. Specht, W. Turner, C. Lisle, R. Thacker. 2013. Arbor: comparative analysis workflows for the tree of life.  PLOS Currents 5.  DOI: 10.1371./currents.tol.099161de5eabdee073fd3d21a44518dc.

Eduardo Ruiz-Sanchez and C.D. Specht. 2013. Influence of the geological history of the Trans-Mexican Volcanic Belt on the diversification of Nolina parviflora (Asparagaceae: Nolinoideae). Journal of Biogeography. 40(7): 1336-1347.  DOI: 10.1111/jli.12073.

Feng-Xia Xu, Dong-Qin Chen, and C.D. Specht. 2013. Comparative microsporogenesis and anther development of selected species of Magnoliaceae.  Nordic Journal of Botany. DOI: 10.1111/j.1756-1051.2012.01445.x

V.S.F.T. Merckx, J. Kissling, H. Hentrich, S.B. Janssens, C.B. Mennes, C.D. Specht and E.F. Smets. 2013. The phylogenetic relationships of the mycoheterotropic genus Voyria and their implications for the early biogeographic history of Gentianaceae.  American Journal of Botany. DOI: 10.3732/ajb.1200330.

V.S.F.T. Merckx, E.F. Smets and C.D. Specht. 2013. Biogeography and conservation of mycoheterotrophic plants. In V. Merckx [ed.], Mycoheterotrophic plants: The Biology of Plants Living on Fungi. Springer Science, New York, NY. 

C.D.Specht, Roxana Yockteng, Ana M.R. Almeida*, B.K. Kirchoff, W.J.Kress. 2012. Homoplasy, Pollination, and emerging complexity during the evolution of floral development in the tropical gingers (Zingiberales). Botanical Review 78: 440-462.

H. Maas-Van de Kamer, P.J.M. Maas and C.D. Specht. 2012. Costus loangensis, an exciting new species from Gabon.  Phytokeys 18: 11-18.

G.V. Cron, C. Pirone**, M.E. Bartlett*, W.J. Kress and C.D. Specht. 2012. Phylogenetic relationships and evolution in the Strelitziaceae (Zingiberales). Systematic Botany 37(3): 606-619. doi: 10.1600/036364412X648562

T. Renner* and C.D. Specht. 2012. Molecular evolution and selection patterns of class I chitinase genes in the carnivorous plants of the Caryophyllales. Molecular Biology and Evolution. doi: 10.1093/molbev/mss106

A. Angelova, Z. Shenghuo, Y. Lin, J. Wong+, and C.D. Specht. Development and deployment of a large-scale flower recognition mobile app. NEC technologies report, 2012.

V.S.F.T. Merckx, S.B. Janssens, N.A.Hynson, C.D. Specht, T.D.Bruns, and E.F. Smets. 2012. Mycoheterotrophic interactions are not limited to a narrow phylogenetic range of arbuscular mycorrhizal fungi. Molecular Ecology, 21(6): 1524-1532. doi: 10.1111/j.1365-294X.2012.05472.x.

T. Renner* and C.D. Specht. 2011. Escaping the trap: Redefining carnivorous plant gland evolution in the Caryophyllales. International Journal of Plant Sciences 172(7): 889-901.

A. Carroll** and C.D. Specht. 2011. Understanding plant cellulose synthases through a comprehensive investigation of the cellulose synthase family sequences. Frontiers in Plant Genetics and Genomics (Front. Plant Sci.) 2:5.

M.E. Bartlett* and C.D. Specht.  2011. Changes in copy number and expression pattern of the TEOSINTE BRANCHED-like genes in the Zingiberales provide a mechanism for evolutionary shifts in symmetry across the order.  American Journal of Botany, 98(2):227-243.

David B. Wake, Marvalee H. Wake and Chelsea D. Specht. 2011. Homoplasy: from detecting pattern to determining process and mechanism of evolution. Science. 331:1032-1034.

A.D. Poulsen and C.D. Specht. 2010. A new species of Costaceae from Borneo. Gardens Bulletin Singapore 62 (1): 143-150. PDF
C. Sass and C.D. Specht. 2010. Phylogenetic estimation of the core Bromeliads with an emphasis on the genus Aechmea (Bromeliaceae). Molecular Phylogenetics and Evolution. 55(2): 559-571. PDF
M.E. Bartlett and C.D. Specht. Evidence for the involvement of GLOBOSA-like gene duplications and expression divergence in the evolution of floral morphology in the Zingiberales. New Phytologist, 187: 521-541. PDF
C.D. Specht and M.E. Bartlett. 2009. Flower evolution: the origin and subsequent diversification of the angiosperm flower. Annual Review of Ecology Evolution and Systematics 40:217-243. PDF
Kirchoff, B.K., L.P. Lagomarsino, W.H. Newman, M.E. Bartlett & C.D. Specht. 2009. Early floral development in Heliconia latispatha and comparative evolution across Zingiberales. American Journal of Botany 96(3):1-15. PDF
Renner, T., J. Bragg, H.E. Driscoll, J. Cho, A. Jackson & C.D. Specht. 2009. Viral Induced Gene Silencing as a tool for investigating floral developmental genetics in the Zingiberales. Molecular Plant 3:1-11. PDF
Ling-Jing Chen, Zhao-Yan Diao, C.D. Specht, and Z. Renee Sung. 2009. Molecular Evolution of VEF-Domain-Containing PcG Genes in Plants. Molecular Plant 2:738-754. doi:10.1093/mp/ssp032
Nguyen, N, H.E. Driscoll, and C.D. Specht. 2008. A molecular phylogeny of the wild onions (Allium; Alliaceae) with a focus on the western North American center of diversity. Molecular Phylogenetics and Evolution 47: 1157-1172 PDF
Bartlett, M.E., B.K. Kirchoff, and C.D. Specht. 2008. Epi-illuminescence microscopy coupled to in situ hybridization and its utility in the study of evolution and development in non-model species. Development, Genes and Evolution 218(5): 273-279. PDF
Kress, W. J., M. Newman, A. Poulsen, and C. D. Specht. 2007. An analysis of generic circumscriptions in tribe Alpinieae (Alpinioideae: Zingiberaceae). The Gardens’ Bulletin Singapore 59: 113-128.
Sass, C., D. Little, D.W. Stevenson and C.D. Specht. 2007. DNA-Barcoding in the Cycadales: Testing the potential of proposed barcoding markers for species identification of Cycads. PloS-ONE. 11: e1154. PDF
Funk, Vicki and C.D. Specht. 2007. Meta-trees: grafting for a global perspective. Proceedings of the Biological Society of Washington. 120(2): 233-241. PDF
Davis, J.I., G. Petersen, O. Seberg, D.W. Stevenson, C.R. Hardy, M.P. Simmons, F.A. Michelangeli, D.H. Goldman, L.M. Campbell, C.D. Specht, and J.I. Cohen. 2006. Are mitochondrial genes useful for the analysis of monocot relationships? Taxon 55:857-870.
Sjölander, K. and Specht, C.D. “Functional prediction through phylogenetic inference and structural classification of proteins.” 2007. In R. Apweiler [ed.], The Encyclopedia of Genetics, Genomics, Proteomics, and Bioinformatics. J. Wiley & Sons, Ltd., West Sussex, UK.
Specht, C.D. “Gondwanan Vicariance or Dispersal in the Tropics? The biogeographic history of the tropical monocot family Costaceae (Zingiberales).” 2006. In J. T. Columbus, E. A. Friar, C. W. Hamilton, J. M. Porter, L. M. Prince, and M. G. Simpson, [eds.], Monocots: comparative biology and evolution. Rancho Santa Ana Botanic Garden, Claremont, California. PDF 277 kb
Kress, W.J. and Specht, C.D. “The evolutionary and biogeographic origin and diversification of the tropical monocot order Zingiberales.” 2006. In J. T. Columbus, E. A. Friar, C. W. Hamilton, J. M. Porter, L. M. Prince, and M. G. Simpson, [eds.], Monocots: comparative biology and evolution. Rancho Santa Ana Botanic Garden, Claremont, California. PDF 277 kb
Petersen, G., O. Seberg, J.I. Davis, D.H. Goldman, D.W. Stevenson, L.M. Campbell, F.A. Michelangeli, C.D. Specht, M.W. Chase, M.F. Fay, J.C. Pires, J.V. Freudenstein, C. R. Hardy and M.P. Simmons. 2006. “Mitochondrial DNA and Monocot Phylogeny.” In J. T. Columbus, E. A. Friar, C. W. Hamilton, J. M. Porter, L. M. Prince, and M. G. Simpson, [eds.], Monocots: comparative biology and evolution. Rancho Santa Ana Botanic Garden, Claremont, California. PDF 380 kb
Specht, C.D. and Stevenson, D.W. 2006 “A revised taxonomy for Costaceae (Zingiberales).” Taxon 51(1): 153-163. PDF 277 kb
Specht, C.D. 2006. “Systematics and Evolution of the tropical monocot family Costaceae (Zingiberales): a multiple dataset approach.” Systematic Botany 31(1): 89-106. PDF 2332 kb
Specht, C.D. 2005. “Phylogenetics, Floral Evolution, and Rapid Radiation in the Tropical Monocot Family Costaceae (Zingiberales).” Pp. 29-60 in A.K. Sharma and A. Sharma, eds. Plant Genome: Biodiversity and Evolution. Science Publishers, Inc. Enfield, NH.
Kress, W. J. and C. D. Specht. 2005. “Between Cancer and Capricorn: phylogeny, evolution, and ecology of the tropical Zingiberales.“ In I. Friis and H. Balslev, eds. Proceedings of a symposium on plant diversity and complexity patterns - local, regional and global dimensions. Biologiske Skrifter, The Royal Danish Academy of Sciences and Letters, Copenhagen.
Davis, J.I., D.W. Stevenson, G. Petersen, O. Seberg, L.M. Campbell, J.V. Freudenstein, D.H. Goldman, C.R. Hardy, F.A. Michelangeli, M.P. Simmons, C.D. Specht, F. Vergara-Silva, and M.A. Gandolfo. 2004. “A phylogeny of the monocots, as inferred from rbcL and atpA sequence variation, and a comparison of methods for calculating jackknife and bootstrap values.” Systematic Botany 29(3): 467-510. PDF 871 kb
Specht, C.D., Kress, W.J., Stevenson, D.W., and DeSalle, R. 2001. “A Molecular Phylogeny of Costaceae (Zingiberales).” Molecular Phylogenetics and Evolution 21(3):333-345. PDF 311 kb
Stevenson, D.W., Davis, J.D., Freudenstein, J.V., Hardy, C.R., Simmons, M.P., and Specht, C.D. 2000. “A Phylogenetic analysis of the Monocotyledons based on Morphological and Molecular character sets, with comments on the placement of Acoris and Hydetellaceae.” Monocots: Systematics and Evolution. Eds K.L. Wilson and D.A. Morrison. CSIRO:Melbourne.
Lim, J., Heliarutta, Y., Specht, C.D., Jung, J., Sims, L., Bruce, W.B., Diehn, S., and Benfey, P.N. 2000. “Molecular Analysis of the SCARECROW Gene in Maize Reveals a Common Basis for Radial Patterning in Diverse Meristems.” The Plant Cell 12:1307-1318. PDF 445 kb
Honors and Awards
Grady Webster Award - 2016
Fellow - California Academy of Sciences - 2015
Kavli Fellow - 2011
Hellman Award - Hellman Family Faculty Fund - 2009
Presidential Chair Fellow - Division of Teaching and Learning - 2009
Prytanean Faculty Award - Prytanean Honors Society Alumni - 2008